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oP BULLETIN

ENTOMOLOGICAL RESEARCH.

ISSUED “BY? TEE “IMPERTAL
BUREAU: Or ENTOMOLOGY.

Hmrror. Lh DERECTOR.

VOL. Xil.

LONDON :
THE IMPERIAL BUREAU OF ENTOMOLOGY,
41, QUEEN’s GATE, S.W.7.
1921-22.

QL&®

IMPERIAL BUREAU OF ENTOMOLOGY.

Bonorary Committee of Danagement.

The EARL BUXTON, G.C.M.G., Chairman.

Lieutenant-Colonel A. W. Atcock, C.I.E., F.R.S., London School of
Tropical Medicine.
Major E. E. Austen, D.S.O., Entomological Department, British
Museum (Natural History).
Dr. A. G. BAGSHAWE, C.M.G., Director, Tropical Diseases Bureau.
Major-General Sir J. Rose BRaprorD, K.C.M.G., C.B., C.B.E., F.RS.,
Medical Adviser to the Colonial Office. :
Major-General Sir Davip Bruce, K.C.B., F.R.S., Chairman of the
Governing Body, Lister Institute.
Mr. J. C. F. FRyYER, Entomologist to the Ministry of Agriculture.
Sir SIDNEY F. HARMER, K.B.E., F.R.S., Director, British Museum
(Natural History).
Professor H. MAXweELL LEFroy, Imperial College of Science and
Technology.
The Hon. Sir E. Lucas, Agent-General for South Australia.
Dr. R. STEWART MacDouGa_t, Lecturer on Agricultural Entomology,
Edinburgh University.
Sir JOHN McFApyYEAN, Principal, Royal Veterinary College, Camden
Town.
Sir DANIEL Morris, K.C.M.G., Late Adviser to the Colonial Office
in Tropical Agriculture.
Professor R. NEWSTEAD, F.R.S., Dutton Memorial Professor of
Medical Entomology, Liverpool University.
Professor G. H. F. NuTTati, F.R.S., Quick Professor of Protozoology,
Cambridge.
Professor E. B. PouLTon, F.R.S., Hope Professor of Zoology, Oxford.
Lieutenant-Colonel Sir DAvip PRAIN, C.M.G., C.I.E., F.R.S.
Sir H. J. ReaD, K.C.M.G., C.B., Colonial Office.
The Honourable N. C. ROTHSCHILD.
Dr. HuGcu Scott, Curator in Entomology, Museum of Zoology,
Cambridge.
Sir ARTHUR E. SHIPLEY, G:B.E., F.R-S., Master of Christ’s: College,
Cambridge.
Mr. R. A. C. SPERLING, C.M.G., Foreign Office.
Sir STEWART STOCKMAN, Chief Veterinary Officer, Ministry of
Agriculture.
Mr. F. V. THEOBALD, South Eastern Agricultural College, Wye.
Mr. C. WarBurTON, Zoologist to the Royal Agricultural Society of
England.
The Chief Entomologist in each of the Self-governing Dominions
is ex officio a member of the Committee.
General Secretary.
Mr. A. C. C. PARKINSON (Colonial Office).
Director and Editor.
Dr. Guy A. K. MARSHALL, C.M.G.

Assistant Director.
Dr. S. A. NEAVE.

Head Office.—British Museum (Natural History), Cromwell Road,
London, S.W.7.
Publication Office—41, Queen’s Gate, London, S.W.7.

(6127) Wt.P6/200 1000 5/22 Harrow G,75.

CONTENTS:

ORIGINAL ARTICLES:

AUSTEN, Major, E. E.
A Contribution to Knowledge of the Blood-sucking Diptera of
Palestine, other than Tabanidae
Some Siamese Tabanidae
BALFrour, Dr. ANDREW.
Mosquito Breeding in Saline Waters
BRUNETTI, E.
A New Pipunculid Parasitic on Leaf-hoppers in India
BRYANT, G. E.
Some New Injurious Phytophaga from Africa
CARTER, HENRY F.
A Revision of the Genus Leptoconops, Skuse
D EL MMEREZ DE CHARMOY, D. & GEBERT, S.
Insect Pests of Various Minor Crops and Iruit Trees in Mauritius
Davipson, Dr. J.
Biological Studies of Aphis rumicrs, Linn.

Dopp, ALAN P.
A New Chalcid Parasite of Euthyrrhinus meditabundus

Dry, F. W.

Flax Caterpillars in Kenya Colony, with Special Reference to the
Limitations of the Roping Method of Combating them ae

The Red Scale, Chryvsomphalus aurantit, Mask., in Kenya Colony ..

The Egg Parasites of the Coffee Bug ( Antestia lineaticollis, Stal) in
Kenya Colony

Trypanosomiasis in the Aieenee a echo and al Elena ieee
possibly carried by Szmulium in Kenya Colony

Epwarps, F. W.
Mosquito Notes ‘
A Revision of the Mosquitos of the Palaearctic ‘Region

PERRIS, (Ga FE.
Some Coccidae from Eastern Asia
GEBERT, S. & D’EMMEREZ DE CHARMOY, D.
Insect Pests of Various Minor Crops and Fruit Trees in Mauritius
GREEN, E. FE. & Laine, F.
Coccidae from the Seychelles
Gee
The Bionomics of Zabanus aprepes and other Australian Tabanidae
The Life-history of Euthyrrhinus meditabundus, Fabr., an eee
Weevil Pest of Mango Trees in Australia
On some Australian Termites of the Genera Drepanotermes, Hami-
termes and Leucotermes
JAMES, LiEuUT.-COLONEL S. P.
Mosquito Life in Surrey during 1921
Laine, F. & GREEN, E. E.
Coccidae from the Seychelles

PAGE

107
431

469

vi CONTENTS.

LaMBoRN, W. A. PAGE
The Nature and Function of the Caudal Tufts of Malayan Anopheline

Pal vac eee a ae ee)

The Mosquitos of some Ports of China and Japan _ ae 401

Ltoyvp. DR EL.
Notes on a Colour Tropism of Astevochiton ( Aleurodes) vaporariorum,

Westwood = a8 is Bs = a a -. | 300
RwEeGe ar
On Three New Species of Indian Braconidae .. as oe si, 129

MACGREGOR, MALco”m E.
The Influence of Drought upon Mosquito Life in Surrey... oo, 205

MARSHALL, Dr. G. A. K.
On New Species of Curculionidae attacking Forest Trees in India .. 165

MOKRZECKI, PROF. S. A.
Agrilus foveicollis, Mars., as a Cause of the Decay of the Culture of

Roses in Bulgaria a an Aa ie hs “es 520 GOO
Morris, HuBert M.
The Larval and Pupal Stages of the Bibionidae. se oc py 4
Patton, Major W. S.
Notes of the Myiasis-producing Diptera of Man and Animals 32259
Notes on the Species of the Genus Musca, Linnaeus. Part I. se LiL
PAWAN, J. EL.
On the Eggs and Ov os of Hee a ( Janthinosoma) posticata,
Wied. (Culicidae) : aes Ss i s. 481

Pomeroy, A. W. J.
New Species of African Simuliidae and Further Studies of the Early

Stages .. : SP a zs it a Ye eee oy
RAMACHANDRA Rao, Y.
Notes on the Life-histories of Two Mesopotamian Moths... .. 477

SCorr, OR. HUGH.
the Ptmid Beetle, eh cake: globulum, Solier, breeding in
Argol se we a se on av a oe. aloe

SUBRAMANIAM, T. V.
Some Natural Enemies of Mango Leaf-hoppers (Jdiocerus spp.) in

India oe vi Ei ee 8 = 28 ae .. 465
Tams, W. H. T.
ee of a New Species of Eprpyrops from South India .. 468
THEOBALD, F.
A New Aphid Genus and Species found in England .. ae “ff A29

Uvarov, Dr. B. P.
A Revision of the Genus Locusta, L. (Pachytylus, Fieb.), with a New
Theory as to the Periodicity and Migrations of Locusts .. av 185
WATERSTON, Dr. JAMES.
On some Bornean Fig-insects (Agaonidae, Hymenoptera Chalcidoidea) — 35

MISCELLANEOUS.
Collections Received .. is Oye eb 105, 203, 361, 483

Vil

Pili.

Breeding-places of Tabanus spp. and Silvius notatius
Mango Tree attacked ie Mee cee meditabundus,

F.
Wings of C ulicoides spp.
Spiracles of Larvae of Myiasis- _producing Flies
Fig. 1—A Typical Case of ‘‘ Simulium Disease ”
Fig. 2.—View of Habitat of Simulium ze
Views of Habitats of Simultum in Kenya Colony
Termitaria of Drepanotermes silvestrit, Hill
Termitaria of Hamitermes perplexus, Hill
Termitarium of Hamitermes meridionalis, Vrogg.
Termitaria of Hamutermes perplexus, Hill

Male Genitalia of West African Species of S¢muliwm ..

Respiratory Filaments of Pupae of Simulium
Parasites of Mango Leaf-hoppers ( /dzocerus spp.)
Eges of Psorophora posticata, Wied.

PAGE

facing 62

66
124
238
232
232
232
400
400
400
400
464
464
470
482

Page

Wis
167,

72,
175,
177,
178,

285,
350
351
Poo)! i
3ol,
SOL;
350,
423,
425,

’

Vili

ERRATA;

line 53, for “ vishuni’’ read “‘ vishnut”’
10 lines from bottom, for “ Dipterocarpus’”’ read “‘ Dipterocarpus
; aE:

tuberculatus

lines 22 and 24, for ‘‘ malabarica’’ read “‘ malabaricum ”

lines 10 and 12, for ‘“‘ Eugenia jaman”’ read “ Eugenia gambolana ’

: Claat: : ” S Z NS =

line 8, delete ‘* Punjab ’

line 25, for “‘ Punjab’”’ read “‘ United Provinces’ and for “ Rani
Range ”’ read “ Rani Rau ”

line 28, for ‘“‘ Drouart ’’ read “‘ Drouard ”’

column 3, line 28, for “ mauritianus ’’ read “ mauritianus ”

column 2: line Bly tor 13255, reads o26; |

column 3, after “ tvitaeniorhynchus ’” add “ turkhudi var. 280”

column 3, line 24, for 325" read, 326.’

column 8, last line for ‘“‘ zammititit’’ read “ zammattii ”

line 2, for “‘ vaporarorium ” read “ vaporariorum ”

last line

s
: : i for “ Johnson’ read “ Johnston’
lines 1, 3, ete: if J z

IMPERIAL BUREAU OF ENTOMOLOGY.

BU EE TIN

ENTOMOLOGICAL RESEARCH.

Vor; XT. 1921.

A EREVISION, (OF + DHE “GENUS LEPTOCONOPS: SKUSE:
By Henry F. Carter,
School of Tropical Medicine, Liverpool.

INTRODUCTION.

The genus Leptoconops was erected in 1890 by the Australian dipterist Skuse for a
minute black Chironomid fly which he captured at Woronora, near Sydney, New
South Wales. This genus he placed in the last of the three sections into which he
divided the family CHTRONOMIDAE, namely, the CERATOPOGONINA. The fly greatly
resembled a small species of Simulium in general facies, and presented certain
peculiar characters which rendered it strikingly distinct from other members of the
section. Indeed, certain authors (Mik 1894, Johannsen 1905) have since drawn
attention to the marked similarity in the wing venation of an American species
(L. torrens, Twns.) and Corynoneura, a genus of the subfamily CHIRONOMINAE. Later
(1907) Noé suggested the formation of an additional subfamily—LEPTOCONOPINAE—for
the Australian and allied species (at that time classified in three genera), and Malloch
(1915), although acknowledging that the genus Tersesthes, Twns. (a synonym of
Leptoconops), was unknown to him, associated it with the CHIRONOMINAE rather than
with the CERATOPOGONINAE. Apart from the wing venation, however, Leptoconops
shows no affinities with Corynoneura, but rather agrees with the Ceratopogonine
midges, particularly in regard to the structure of the thorax and mouth-parts.
Several species of Leptoconops have now been described, but owing to differences in
the interpretation of, or slight variations in, some of the generic characters given by
Skuse, as well as to subsequent descriptive errors, they have, in greater part, been
referred to the genera Tersesthes, Twns., and Mycterotypus, Noé. As was to be expected,
however, the close agreement exhibited in the diagnoses of these genera and of
Leptoconops has caused several authors to suggest their identity; but indefinite
or partial conclusions* only were reached, since the genotype of Leptoconops (L.
stygius, Sk.) was not re-examined.

Through the courtesy of several gentlemen (individual acknowledgments are
made in the text of the systematic portion of this paper) I have been privileged

* Castellani and Chalmers (1913 and 1919) give Tersesthes and Mycterotypus as synonyms of
Leptoconops, stating that in so doing they follow Kieffer. The latter author, however, did not
definitely reject these names, and in his recent works still refers to them.

(2416) Wt.P8/170 1,000 5/21 Harrow G75. A

2 HENRY F. CARTER.

to examine the majority of the known species of this genus, including L. stygius,
and have thus been able to decide certain questionable morphological characters
which were largely responsible for the previous confusion. The new species
described herein were received from the Imperial Bureau of Entomology (per
Dr. G. A. K. Marshall), the British Museum (per Mr. F. W. Edwards), and the
United States National Museum (per Prof. L. O. Howard and Dr. J. M. Aldrich) ;
the types and co-types of these species have been returned to the collections
from which they came, as subsequently indicated at the end of the respective
descriptions.
SYNONYMY AND CLASSIFICATION.

Skuse’s definition of Leptoconops is as follows :—‘‘ Antennae in the female 2 +
11-jointed ; first joint of scapus large, disciform ; second smaller, globose ; flagellar
joints globose, gradually diminishing in size, more ovate towards the apex, terminal
joint elongate-ovate. Proboscis prominent. Palpi four-jointed ; first and second
joints small, third greatly incrassated, about three times the length of the first or
second: fourth not as long as the last, slender, cylindrical. Wings naked. All
longitudinal veins taking their origin at the base of the wing. Marginal cross-vein
present. Middle cross-vein wanting. Fourth and fifth longitudinal veins only
forked.”

In 1893 Townsend founded the genus Tersesthes, but although acquainted with
Skuse’s work, would seem not to have appreciated the close resemblance between
his genus and Leptoconops. Practically the only points of difference between the
two genera that can be obtained from his description are that the palpi are composed
of three segments and the wings covered with microscopic hairs ; but in his figure
of the wing the costa is extended to the apex, terminating near the upper branch
of the fourth vein.

Noé (1905) erected the genus Mycterotypus for two Italian midges. He was,
however, doubtful as to its distinctiveness from Leptoconops and Tersesthes, and
was unable to decide whether the differences between these two genera and
Mycterotypus were real or apparent. But in view of the facts that he could
distinguish satisfactorily only three palpal segments, that the venation apparently
differed in several particulars, and chiefly that he believed the “ cerci”’ (lamellae)
to be absent in Leptoconops, he finally resolved to place his species in a separate
genus.

Johannsen (1905) concluded that Leptoconops and Tersesthes were very closely
related, if not identical, and that they could only be distinguished by the segmentation
of the palpi. Kieffer (1906), however, retained all three of these genera, but subse-
quently (1908) suggested that they were probably synonymous and that Townsend’s
figure of the wing showing the extension of the costa to the apex was inaccurate.
Langeron (1913), after studying the venation of a Tunisian species and the figures
of the wings of Leptoconops and Tersesthes, considered the former genus and
Mycterotypus very nearly allied, but expressed surprise that Kieffer should think
Tersesthes and the latter identical. Lutz (1913) was the first author who actually
compared specimens of any of these genera; he examined both of Noé’s species of
Mycterotypus and Townsend’s Tersesthes torrens, and definitely decided that they
were congeneric. De Meijere (1915) briefly discussed the differences which existed,
or were said to exist, between the three genera, and concluded by adopting Kieffer’s
suggestion of identity and listing all the species then known under Leptoconops. The
latter author, however, does not appear to have reached a definite decision in this
regard, for although placing Tersesthes and Mycterotypus under Leptoconops, he
yet (1917 and 1918) retains the names to indicate groups of species ; moreover, in
spite of previous suggestions of inaccuracy, he (1917) still accepts Townsend’s
interpretation of the wing venation, but employs it for purposes of specific
differentiation !

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 3

Comparisons of the genotype species of Leptoconops (L. stygius), Tersesthes (T.
torrens) and Mycterotypus (M. bezzi1) have shown definitely that all are congeneric,
and therefore the last two generic names must sink under Leptoconops. The value
and nature of the various characters upon which the separation of these genera was
based will be discussed later in connection with the external morphology.

In a supplement to his 1918 paper Kieffer divided Leptoconops into three genera
as follows :—

“1. Antennes de la 2 composées de 12 articles ; crochets tarsaux simple, égaux,
petits; type: kertesz1, Kieff. (Région palearctique).
Genre Holoconops, n.g.
Antennes de la 2 composées de 13 articles

“2. Crochets tarsaux de la 9 bifides, les 2 rameaux inégaux (Indes orientales).
Genre Schizoconops, n.g.

Crochets tarsaux de la Q simple, inégaux au tarse antérieur, égaux au
tarse postérieur (Australie). Genre Leptoconops, Skuse.”

This classification cannot be maintained for two reasons: first, the two types of
differential characters used are not of the same value and are neither of them sufficient
for purposes of generic separation, and, secondly, the description of the claws of
Leptoconops (with type L. stygius) is erroneous. Holoconops may be retained with
advantage as a subgenus by reason of the antennal structure of the females, Lepto-
conops, in the restricted sense, being reserved for those species with the full comple-
ment of segments. Lepftoconops (sens. str.) can, if necessary, be further separated
into two groups on the structure of the claws ; but even if such groups be worthy of
subgeneric rank, the name Schizoconops cannot be employed, since the genotype of
Mycterotypus (M. bezzit was the first species described by Noé) possesses toothed
claws, thus giving the latter name priority.

In 1915 de Meijere described a species (L. albiventris) from New Guinea which
differed from all other Leptoconops in the structure of the ovipositor. The discovery
of an African species possessing a similar ovipositor, and the fact that in both species
this character is supported by others not present in Leptoconops, appear to justify
the erection of a new genus ; this is described on p. 24 under the name Acanthoconops.

DISTRIBUTION.

The members of the genus Leptoconops (sens. lat.) are widely distributed, but so
far as is known are confined to countries lying approximately between the parallels of
latitude 40° North and 35° South. Representatives occur in Italy, Sardinia, Asia
Minor, Bengal, Siam, Northern, Central and South-West Africa, United States of
America, Cuba, Brazil and Australia. The two species of Acanthoconops are found in
New Guinea and Zanzibar.

BIOLOGY.

Nothing is known of the life-histories or early stages of these flies, and compara-
tively little concerning the habits of the adults. In fact, the only species which have
been studied in any detail are L. bezzii and L. irritans, both of which were investigated
by Noé (1905 and 1907) in the Roman Campagna; and to a less extent L. kerteszi
(Mycterotypus laurae), observations on which have been recorded by Weiss (1912) and
Langeron (1913). Certain authors (Noé, Lutz, and, according to Weiss, Bezzi) have
suggested, in view of the greatly developed ovipositor, that the eggs are not deposited
superficially, and that the larvae are terrestrial, living among the roots of grasses, etc.
Noé, indeed, excluded water as the larval habitat, since he failed to rear either of the
Italian species from various aquatic larvae collected during his investigations, even
although adults of both flies were present in enormous numbers at the time ; and be-
cause both species appeared and subsequently increased greatly in numbers at a period

(2416) A2

4 HENRY F. CARTER.

when water was relatively scarce. Weiss, on the other hand, states that at Tabeditt,
South Tunis, L. kerteszi and Simulium maculatum, Mg. (S. lineatum, Fries) occur in
common swarms, and show such close association as adults that he believes the
immature stages of the two flies will be found near together. In this connection I
am indebted to Dr. J. M. Aldrich, of the United States National Museum, for the
following interesting observation, which indicates that the larvae of L. torrens, at
least, are terrestrial in habit. He writes: “ Several years ago, before my connection
with the Museum, I identified a few specimens as Tersesthes torrens. The two females,
which I retained in my private collection, have labels reading, ‘Maxwell, N.M. Reared
from pupae of Tachina mella, Webster. No. 11154. C.K.Wildermuth.’ The
significance of the rearing record is that the adults made their appearance in a breeding
cage containing pupae of the fly ; this proves not parasitism, but that the larvae are
terrestrial, as is known to be the case in some of the CERATOPOGONINAE.”’

A perusal of the data subsequently given in the systematic portion of this paper
in connection with the habitats of the different species provides some indication of
the types of country in which these flies are known to occur, and of the avidity with
which they bite. It will be seen that they have been found in what appear to be
both relatively dry and well-watered districts, also in low-lying as well as mountainous
regions. Noé states that L. ivritans is especially prevalent in the coastal plains of
the Roman Campagna. Langeron records L. kerteszi from marshy places in the desert
in southern Tunis ; Willcocks (1918) has observed the same insect in the Nile delta ;
while Weiss, Townsend (1893) and others have found species at altitudes varying
from 1,600 ft. to 7,000 ft. So far as is known, they are diurnal insects, continuing
their activities throughout the hottest parts of the day. Noé states that in Italy
L. bezzu and L. trritans (locally known as ‘ serapiche’) appear towards the end of
May, reach their maximum abundance about the middle of June, and then gradually
decrease in numbers and disappear early in September. The females of several species
are known to bite man and his domestic animals, and to the former at least sometimes
cause great annoyance and inconvenience. In some districts they have gained an evil
reputation, and indeed at certain seasons would appear to be very serious pests, since
at the period of maximum abundance the females of some species attack in dense
swarms. According to Noé, L. irvritans may occur in such swarms that no defence
is possible, and he adds that labourers working on the railway from Rome to Pisa
were sometimes forced to take shelter in order to escape these massed attacks. L.
kerteszt is also known to adopt this habit (Weiss), and Sambon (1913) states that
Chalmers in the Nile delta “ obtained from the fellahin a very definite history of small
black, blood-sucking flies appearing in swarms, and Dr. Gough told him that this fly
scourge of the delta is undoubtedly Leptoconops ;”” Chalmers was unable to find
Simulium in this region, but L. kerteszi is abundant in certain localities (Willcocks).
Pratt (1907), writing on North American blood-sucking midges, says that Barber
considers Leptoconops (Tersesthes) ‘“‘ much worse asa pest than any Ceratopogon (sens.
lat.) he has ever encountered.” The bites are painful and the subsequent local
reactions irritating and persistent ; L. zvritans, in addition, also causes considerable
discomfort by crawling about the body, among the hair, beneath the clothes, in the
ears, etc. (Noé). Noé’s observations, however, tend to indicate that in some
respects the habits of different species are not identical ; L. bezzi1 is said to prefer the
neighbourhood of houses and outbuildings rather than the open plains, and to be
particularly attracted by poultry, in the houses of which the females shelter over-
night. Further, although the females apparently prefer blood for food, Noé has seen
both sexes of this species on flowers, and remarks that the males are especially fond
of Euonymus.

ASSOCIATION WITH DISEASE.

Grassi (1901) endeavoured to obtain the experimental infection of L. ivritans, Noé,
with malaria parasites. He fed twenty-eight ‘“‘ wild’’ females of this species upon

A REVISION OF THE GENUS LEPTOCONO@PS, SKUSE. 5

three patients, two of whom were infected with Plasmodium falciparum and one with
Plasmodium falciparum and Plasmodium vivax. No infection of these midges occurred
although females of Anopheles maculipennis fed at the same time and on the same
patients became infected from two of the three cases. Grassi concluded that
Leptoconops (Centrotypus) was not able to transmit human malaria.

In 1913 Sambon extended the possible carriers of pellagra to include, besides the
SIMULIIDAE, certain blood-sucking midges; he particularly referred to Leptoconops

in this connection, basing his chief reason for so doing on Chalmers’s observations (see
above) made in pellagra districts in Lower Egypt.

Chatton and Blanc (1917) in a paper on Toxoplasma and toxoplasmosis of the
gundi (Ctenodactylus gundt) discuss the natural ectoparasites of this animal; among
these they make specific mention of two biting flies, namely L. kerteszi, Kieff.
(Mycterotypus laurae, Weiss), and Simulium maculatum, Mg. (lineatum, Fries).

SYSTEMATIC ACCOUNT.
Genus Leptoconops, Skuse (sens. lat.).
Leptoconops, Skuse, Proc. Linn. Soc. N.S.W. (2) iv, p. 288 (1890).
Tersesthes, Twns., Psyche, vi, p. 369 (1893).

Centrotypus, Grassi (nomen nudum), ‘‘ Die Malaria: Studien eines Zoologen,”’ Jena,
pa LLS (1901):

Mycterotypus, Noé, Atti Accad. Lincei, Ser. 5, Rendiconti, xiv, p. 114 (1905) ;
Arch. Zool. Napoli, ii, p. 101 (1907).

Mycteromyia, Lutz (nec Phil.), Mem. Inst. Osw. Cruz, iv, p. 24 (1912); ibid. v,
p. 69 (1913).

Holoconops, Kieff., Ann. Mus. Nat. Hung., xvi, p. 135 (1918).
Schizoconops, Kieff., Ann. Mus. Nat. Hung., xvi, p. 135 (1918).

The external morphology of the adults of this genus has been discussed in detail by
Townsend (1893), Noé (1905 and 1907), and Langeron (1913) in relation to the species
described by them. Owing probably to lack of material, however, no general account
based on the examination of a number of species has yet been given, and it is there-
fore thought advisable—especially in view of the discrepancies which have appeared—
to include here a description of the more important structures.

Fig. 1. Head of : (a) Acanthoconops spinosifrons, sp. n., 9; (b) Leptoconops kertesat var.
americanus, n. (x 90 circa).

6 HENRY F. CARTER.

Head. Eyes bare, widely separate (the space between them varying from one-fifth
to almost one-half the width of the head) in both sexes.* Vertex and occiput bearing
4 few short hairs or bristles ; frons nude or with at most a single pair of short bristles
between the eyes (fig. 1, 6). Clypeus moderately pronounced, rounded, hairy.

Proboscis as long as, or rather less in length than, the height of the head. Mouth-
parts in the female as follows : labium soft and hairy, broad, the labella relatively
large ; labrum strongly chitinised, broad at the base, gradually tapering towards a
rounded apex, the extremity armed with three recurved teeth ; hypopharynx less
strongly chitinised than, but somewhat similar in shape to, the labrum, the apex
devoid of teeth, pointed and scoop-like ; mandibles and maxillaey well-developed,
the former moderately chitinised, relatively broad and obliquely truncate or curved
distally, bearing twelve to twenty-four small, closely apposed teeth, the maxillae
narrower, slightly shorter and more pointed, armed with from twelve to thirty larger
and more widely separated teeth. Mouth-parts in the male less strongly chitinised
than those of the female; extremity of the labrum hairy, mandibles not visible
(? absent) in the single specimen available, maxillae slender, thinly chitinised,

pointed, without teeth.

Fig. 2. Palpiof 9 9 of: (a) L. stygius, Sk.; (6) L. torvens, Twns. ; (c) L. rvhodesiensis, sp. n. ;
(d) L. kerteszi var. americanus, n.; (e) A. spinosifrons, sp.n.; (f) L. siamensis, sp. n. (x 220 circa).

Palpi (fig. 2) composed of four segments. In the female the first and second
segments are reduced in size and often indistinctly separated one from another, the
third incrassate, the last relatively large, representing the combined small fourth and
fifth segments of other CERATOPOGONINAE ; the sensory pit of the third segment is
highly developed, the orifice often large, occupying the greater portion of the inner
aspect, or occasionally small, sub-circular ; terminal segment with an apical whorl of
hairs. In the male the palpi are longer than in the female, and the third segment is
not, or scarcely, swollen.

* In this account remarks concerning male characters are based upon the examination of a
specimen provisionally referred to L. torrens (see p. 15) and Noé’s description of L. bezztt. The
male of L. kerteszi (M. laurae, Weiss) is not considered here, since, as shown on p. 22, its structure
in certain respects is so peculiar that confirmation of Weiss’s observations is desirable before any
definite statements can be made.

+ Langeron (1913) in his description and figure of the mouth-parts of the female of L. kerteszi
var. peneti (M. laurae var. peneti) has, through incorrect interpretation, transposed the names of
these structures.

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 7

The confusion which has arisen in regard to the segmentation of the palpi has been
entirely due to the somewhat rudimentary nature of the first and second segments,
and to the varying degrees of differentiation exhibited by them in certain species.
The palpi have thus been described as possessing four, three, or even two segments.
when the basal ones were overlooked or ignored.

NIC
=,

Fig. 3. Proximal portion of antenna of Acanthoconops albiventris, Meij.,
Q, showing the reduced first segment : side view (x 510 circa).

Antennae set in large subcircular, pale-coloured, thiniy chitinised, depressed areas.
In the female the antennae (fig. 5, a—h, and cf. fig. 1) are pilose, composed of thirteen
(subgen. Holoconops) or fourteen (Leptoconops, sens. str.) segments; first segment
(cf. fig. 3) (ignored by all previous authors except Lutz) cup-like, broad at the apex,
where there is a rim of thickened chitin, narrowing rapidly to a conical base ; second
segment large and rounded, the inter-segmental membrane connecting it to the
chitinised rim of the first, extensive ; segments of the flagellum eleven or twelve in
number, all but the last with a distinct whorl of hairs and two or more transparent
spines; third segment (first of the flagellum) relatively large, pyriform; terminal
segment elongate-oval, considerably longer than any of the others; intermediate
segments (4 to 12 or 13) almost uniform in shape and size, often transversely oval or
subspherical, rarely narrowly oval. In the male the antennae are longer and plumose,
composed of fifteen segments ; the two basal segments similar to those of the female
but larger, the second very large; segments of the flagellum as shown in fig. 4, h,
and described on pp. 16 and 17.

Thorax arched anteriorly but not projecting over the head, with short and some-
what sparsely arranged hairs on the dorsal surface. Situated laterally, near the
anterior margin, and centrally, in front of the scutellum, are small depressions some-
what similar to those found in Culicoides ; in each anterior depression, however, are
two or three separate oval or rounded, seemingly membranous, areas (which frequently
appear as small shining black spots in dry specimens), not a single slit-like area as in
Culicoides. Scutellum with the posterior margin gently rounded, slightly produced
laterally, bearing two or three pairs of strong, and sometimes a few pairs of smaller,
bristles. Post-scutellum strongly arched, nude.

Wings white, iridescent, with the surface entirely covered with minute upright
setae and with a fringe of longer hairs on the distal and posterior margins. The
venation (fig. 7) is characteristic, but the veins, particularly the anterior ones, are ill-
defined and difficult to distinguish. They can best be followed in dry specimens, but
careful manipulation is necessary, as the detection of certain details depends largely
on the angle of view and the illumination ; in mounted specimens (unless stained) the
venation cannot be observed. The costa is short and, in the female. usually terminates

8 HENRY F. CARTER.

well before the middle of the wing. The sub-costa is chitinised and more clearly
defined than any other vein ; it is closely apposed to the first longitudinal vein, and
owing to the folding of the wing surface in this region, frequently obscures, or partly
obscures, the base of the latter. The first and third longitudinal veins (the second is
absent) are separate basally, but fuse distally, forming with the extremity of the costa
a large, slightly raised, yellowish or pale brown area. These two veins are approxi-
mated throughout the greater portion of their course, but diverge slightly before
fusing with one another and the costa; this approximation is usually greatest im-
mediately before the divergence mentioned above, and in several species is so close
that amalgamation has taken place, with the results that an apparent cross-vein has
been formed and a small cell isolated (cf. fig. 8, a-l). The anterior or radio-medial
cross-vein is absent. The general course of the fourth, fifth and sixth veins shows
little variation ; the upper branch of the fourth vein joins the apex of the wing below
the middle, the lower branch is often very indistinct, with an apparently considerable
portion of its base, and sometimes its apex, obsolete. At least three vein-like folds
(indicated in fig. 7 by dotted lines) are present, and of these the most anterior—which
is evidently homologous with the fold just above the upper branch of the fourth vein
in other CERATOPOGONINAE—is relatively strong and conspicuous ; it is, indeed, as
strongly marked as any of the veins except the sub-costa, and by most authors has
been interpreted as the third longitudinal vein.

. (?) torrens, Twns., @: a, clasper; b, claws of front leg ; h, antenna (flagellum).
. kerteszt var. americanus, n., : c, claws of hind leg (near claw foreshortened).

. bez2i, Noé, ¢:d, claws of middle leg.

stygius, Sk., 9: e, claws of front leg (ventral view empodium omitted).
stamensts, sp.n., ©: f, one claw of front leg.

. brasiliensis, Lutz, fe : g, first tarsal segment of front leg.

ax 770); (b,:¢, f * A905 d x 2205 &x 450 ei 260) 7905)

[be
Ee
IE
IB.
he.
iL

—

As will be gathered from a perusal of the subsequent specific descriptions and from
the illustrations referred to above, this account of the wing venation is based upon
the study of the wings of females of several species. Among these are included most
of the forms previously referred to Tersesthes and Mycterotypus, and it therefore
follows that the venation in these insects is in no way peculiar, and that venational
characters which have been advanced for the retention or differentiation of these
genera are either of little value or are the results of incorrect observations.

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 9

Legs moderately long, the hind pair longest, clothed with short hairs. Femora
unarmed. Tibiae each armed distally with a short, stout, ventral spur, those of the
fore and hind legs in addition with one or two oblique rows of bristles. First tarsal
segment of the fore and middle legs about twice the length of the second, of the hind
legs about one and one-half times the length of the second; second to fourth
tarsal segments cylindrical, decreasing in length progressively, the fifth segment
distinctly longer (in L. Jlactetpennis Kieffer states that it is shorter) than
the fourth. Differentiation of the apical bristles of the first and second tarsal

Fig. 5. Terminal segments of antennae of females of: (a) L. siamensis, sp.n.; (6) L. stygius,
Sk.; (c) L. grandis, sp.n.; (ad) L. longicornis, sp. n.; (e) L. rhodesiensis, sp.n.; (f) Acanthoconops
spinosifrons, sp. n.; (g) L. kerteszt var. americanus, n.; (h) L. tovrens, Twns. {x 260 circa).

segments into spines has taken place in most species, and of some of the
ventral bristles of the first segment (fig. 4, g) in a few species. In the latter
case considerable prominence has been given to this character by some writers
in their specific descriptions, and the exact number and arrangement of such spines
has been recorded ; but variation in details is frequent, and may occur not only in

10 HENRY F. CARTER.

different individuals of the same species, but on different legs of the same individual
(cf. footnote p. 20). Claws equal, small, less than one-half the length of the fifth
segment ; in the female similar on all the legs, either simple, each with a bristle arising
from the base (fig. 4,c and ¢) or dentate, with a strong basal tooth (fig. 4, d and‘f) ; in the
male (fig. 4, 0) dissimilar on the fore and middle legs, one simple, the other with a long
basal tooth (in L. bezzit both are said to be dentate), on the hind legs similar, simple.
Empodium in the form of a minute branched bristle.

Abdomen of the female composed of nine segments—the ninth greatly reduced in
size—clothed with short hairs, and bearing distally two exceedingly long, narrowly
conical lamellae (fig. 6). Genital orifice, “anteriorly, with a semi-circular chitinous
border from which arise numerous small and a few long hairs, the latter directed
inwards and backwards over the aperture. Spermathecae usually two in number
(occasionally, e.g., L. kerteszi,a small third spermatheca is present), subspherical or
oval and but slightly produced posteriorly before the junction with the duct. Abdomen
of the male more slender than that of the female, the hypopygium conspicuous.

Subgenus Leptoconops (Skuse), Carter.

As here restricted, this subgenus includes only those species in which the
antennae of the female are composed of fourteen segments; in this sex the species
fall into two groups according to the claws being simple or dentate.

Leptoconops stygius, Skuse.
Leptoconops stygius, Skuse, Proc. Linn. Soc. New South Wales, (2) iv, p. 288 (1890).

Leptoconops skusii, Noé (errer in explanation of Plate v), Arch. Zool. Napoli, i
(1907).

Skuse’s description of this species, the type of the genus Leptoconops, is as follows:—

‘““ 0.—Length of antennae, 0:42 mm. ; expanse of wings, 1-27 mm. ; size of body,
1:66 mm. Entirely black. Joints of antennae with dense light-greyish verticils.
Head and thorax levigate, with minute black hairs. Abdomen about twice the
length of the thorax, opaque, with some minute black hairs; lamellae very long,
slender. Legs slender. Hind metatarsus one-third longer than the second tarsal
joint. In the fore legs, the tibiae rather more than twice the length of the metatarsus.
Wings hyaline, rather weakly iridescent ; costal and first two longitudinal veins
greyish-brownish, the rest pale and indistinct. Auxiliary vein not distinguishable,
apparently wanting; first and second [?.e., third} longitudinal veins reaching costa
before the middle of the anterior border, coment at the tips, tip of second longitu-
dinal vein almost opposite but immediately beyond the tip of the posterior branch of
the fifth longitudinal vein ; marginal cross-vein indistinct ; marginal cell small ; third
longitudinal vein [2.¢., the strong fold situated in the upper portion of the wing]
arcuated, not quite reaching the margin, terminating a little above the apex of the
wing ; fourth longitudinal vein bellied downwards at the middle, reaching the margin
a little below the apex of the wing, the posterior branch detached ; fork of fifth
longitudinal vein wide, the anterior branch twice the length of the posterior.

‘““ Habitat. Woronora (Skuse). October.”

Through the kindness of Professor S. J. Johnston, of Sydney University, I have
been able to examine one of the three specimens of L. stvgius contained in the Macleay
Museum, Sydney, and am thus in a position to supplement the above description with
some important morphological details.

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 1]

Q. —Length of body (specimen mounted in balsam), 2°55 mm.; length of wing,
13mm. ; length of antenna, 0'48 mm. ; width of head, 0°40 mm.

ead. Eyes relatively narrowly Le eeatedl the space between them almost one-
fifth the greatest width of the head; clypeus with two pairs of short hairs. Third
palpal segment (fig. 2, a) relatively not very strongly swollen, elongate, the orifice of
the sensory pore extending over the greater portion of the inner side ; fourth segment
subcylindrical, about two-thirds the length of the third. Antennae (fig. 5, 6): fourth
to thirteenth segments spherical, with the hairs arranged in oblique whorls, and the
spines unequal in size and asymmetrically arranged; terminal segment short,
approximately one and two-thirds as long as broad. Wings with anterior veins as
shown in fig. 8, a.* Legs: first and second tarsal segments without differentiated
spines, except distally. Claws (fig. 4, e) simple and equal, each with a bristle arising
from the base. Lamellae bluntly rounded distally, 0°22 the length of the wing.
Spermathecae two, heavily chitinised, subspherical (diameter 30u) ; the commence-
ment of the duct chitinised for a short distance.

Leptoconops longicornis, sp. nov.

2.—Length of body (two specimens), 3°5 mm. ; length of wing, 2:2 mm. ; length
of antennae, 0°84 mm. ; width of head, 0:44 mm.

Head black,+ the antennal depressions pale brown, sparsely clothed with short
hairs on the occiput and vertex ; clypeus blackish, with three pairs of dark brown
hairs ; eyes not very widely separated, the space between them being approximately
one-fifth the width of the head ; frons with a pair of short hairs situated, one on each
side, near the lower margins of the eyes. Proboscis blackish brown. Palpi blackish
brown, with dark hairs; third segment elongate, relatively slightly swollen, the
orifice of the sensory pit large, occupying the distal two-thirds of the inner side ;
fourth segment subcylindr ical, stout and short, not more than two-thirds the length
of the third. Antennae (fig. 5, d) long, dark brown, bearing short brown hairs, and
long, slightly curved, clear, pointed spines ; fourth to thirteenth segments subspherical
to narrowly oval, from 1-0 to 2:1 as long as broad ; fourteenth segment approximately
six and one-half times as long as broad, equal in length to the three preceding segments
together. Thorax shining black (from indications still existing in the dried specimens
the scutum was probably dark grey pollinose originally), sparsely clothed with short
dark hairs; scutellum normally with three pairs of stout black bristles; pleurae
and pectus shining black. Wings white, iridescent, the fusion of the extremities of the
anterior veins forming an elongate brown spot near the middle of the upper margin ;
anterior veins as shown (fig. 8, 0), the distal interspace clearly defined, fifth vein
bifurcating appreciably before the extremity of the costa. Halteres greyish buff, the
stems somewhat infuscated. Legs uniformly dark brown, clothed with dark hairs ;
tarsi without distinct spines (except perhaps a distal pair on the metatarsi), but with
some of the ventral bristles on the first and second segments stout and _spine-
like. Claws, equal and simple, similar to those of L. stygius. Abdomen dark brown,
with short dark hairs. Lamellae (fig. 6) paler brown, 0°18 the length of the wing.
Spermathecae two, heavily chitinised, subspherical (diameter 50u) ; the origin of
the duct only chitinised.

* I have been totally unable to distinguish the small ‘‘ vein ’’ (termed by Skuse the “‘ marginal
cross-vein ’’) connecting the distal portions of the first and third veins. The first vein, however,
bends sharply upwards just before its termination and at the angle is distinctly swollen, causing
the lower edge of the vein to approach more closely that portion of the third vein immediately
below it ; at. first sight, therefore, the first and third veins appear to be connected at this point
and to enclose a minute distal interspace.

+ The colours given in the descriptions of this and the following species are as observed in
specimens which had been dried after preservation in formalin.

1 HENRY F. CARTER.

Habitat. Interior of Western Australia: (J. W. Dakin), 1915. Five females
(two cotypes) in the British Museum Collection.

Professor Dakin states that these midges do not trouble one before 10 a.m., and
that they disappear at dusk ; in between these hours they bite furiously, and the
bite irritates for days afterwards.

Ree \ ij
5 AM VASES

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rn
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Near ho

Li AG ET EE EOS
hdr bal ee 7).

PAPAL A A Aaat
7.

“777

GAG
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L134

FP
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3
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47
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SASS

CAN

Fig. 6. Leptoconops longicornis, sp. n., ventral view of

extremity of abdomen of 9; g.o., genital orifice; viii,

sternite of eighth segment; ix, sternite of ninth segment ;
a, anus; 6, lamellae; (x 180 circa).

Leptoconops grandis, sp. nov.

?.—Length of body (one specimen), 35mm. ; length of wing, 20 mm. ; length of
antennae, 0°62 mm. ; width of head, 0°44 mm.

Two females of this species, captured at the same place and time and bearing the
same data attached to the label as the specimens of L. longicornis, were included in
the material collected by Professor Dakin. Indeed the two forms were contained in
the same tube and, except in antennal structure and certain details of minor
importance, resemble one another so closely that, in the absence of males, it is difficult

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 13

to know exactly what value to place upon the differences observed. The antennal
structure, however, differs so strikingly that I believe specific separation to be
warranted ; especially since no tendency to intermediate characters occurred in the
small series of specimens obtained, and no obvious variation has been seen in the
antennal structure of species of which numerous examples have been examined.

The antennae (fig. 5, c) are very distinctly shorter than in the preceding species,
and the intermediate segments of the flagellum (4 to 13) are subspherical, being from
1-0 to I'l times as long as broad ; the terminal segment is almost three and one-half
times as long as broad, and is equal in length to the preceding two and one-half
segments together. Minor differences appear to exist in regard to the venation
(cf. fig. 8, 6 and c), and the dark grey pollinosity of the head and thorax ; but, as
indicated above, the appearance of the latter may be that normally found in
L. longicornis.

The two females (cotypes) of this species are in the British Museum Collection.

The three Australian representatives of Leptoconops (L. stygius, L. longicornis and
L. grandis) at present known are closely related, and as a group are characterised by
the relatively narrow space separating the eyes, the structure of the palpi (i.e. in
regard to the relative lengths of the third and fourth segments), and the absence of
spines on the first and second tarsal segments.

Leptoconops braziliensis, Lutz.

Tersesthes braziliensis, Lutz, Mem. Inst. Oswaldo Cruz, v, p. 66 (1913).

The description of this species given below is drawn up from that published by
Dr. Lutz, and from microscopical preparations kindly lent me by him.

2.—Length of body (two specimens), 1°5 mm. ; length of wing, 0'8 mm. ; length
of antennae, 0°32 mm.; width of head, 0:23 mm.

Head dark brown ; clypeus with a few short, dark hairs. Proboscis and palpi
dark brown ; the latter with the third segment moderately swollen and the orifice of
the sensory pit large and oval, the fourth segment subcylindrical; almost equal in
length to the third. Antennae dark brown, with dark hairs and short, slightly curved,
clear spines ; fourth to thirteenth segments transversely oval, from 0°6 to 0°8 as long
as broad ; thirteenth segment somewhat pointed distally, two and one-half times as
long as broad, almost equal in length to the preceding four segments together. Thorax
dark brown, clothed with short hairs ; pleurae and pectus rather paler in colour than
the dorsum. Wangs white, the basal part of the costa waxen yellow-brown, the costa
extending to the middle of the anterior border ; anterior veins arranged as in fig. 8, g,
the fifth vein bifurcating before the extremities of the third vein and costa. Halteres
with pale knobs and brown stems. Legs brown, the tarsi paler; metatarsi of the
four anterior legs (fig. 4, g) with several pairs of distinct spines, of the hind legs with
short stout bristles, but with spines only at the apex (distal pair) ; second tarsal
segments of all the legs with a pair of spines at the apex. Claws simple and equal,
each with a bristle arising from the base. Abdomen dark brown dorsally, the fore and
hind margins of the tergites narrowly paler ; venter pale brown. Lamellae waxen
brown, darker at the extreme base, relatively long, 0°33 the length of the wing.
Spermathecae two, strongly chitinised, oval (334 x 244).

Habitat. Brazil: lower reaches of the Rio Tocantin.

According to Lutz this species sucks blood and often attacks man. It may be
readily distinguished from other species of Leptoconops (sens. stv.) which possess
simple claws by the unusually long lamellae.

14 HENRY F. CARTER.

Leptoconops irritans, Noe.

Mycterotypus irritans, Noé, Atti R. Accad. Lincei, Ser. 5, Rendiconti xiv,
p. 118 (1905) ; Arch. Zool. Napoli, ii p. 138 (1907).

Centrotypus trritans, Grassi (nomen nudum), “Die Malaria: Studien eines
Zoologen,”’ Jena, pp. 118-122 (1901).

This species occurs with L. bezzi1 (see page 17) in the Roman Campagna, where,
according to Noé, it is very abundant from June to the end of July, and may be found
in diminished numbers late in August. The female only is known, and in this sex the
species may readily be distinguished from L. bezzi7 by its general facies. Noé confined
himself to a comparative description, and gave the following principal differences
between it and the latter species.

Size smaller (length, 15 mm.; spread of wings, 2 mm.) ; proboscis and palpi
relatively longer, the former more slender, cylindrical ; antennal hairs sparser and
more spinose ; claws simple, the large basal tooth replaced by a robust bristle ;
abdomen white, becoming isabella-coloured dorsally.

To these I am able to add further details of specific importance, obtained from
Sardinian specimens sent me by Professor M. Bezzi.

Q.—Length of body (one specimen), 1°7 mm. ; length of wing, 1:1 mm. ; length
of antenna, 0°53 mm. ; width of head, 0°30 mm.

Eyes separated by about one-third the width of the head. Antennal segments
4 to 12 transversely oval to spherical, the length from 0°7 to 1:0 the width; terminal
segment about two and one-third times as long as wide, slightly longer than the two
preceding segments together. Scutellum with two pairs of bristles. Metatarsi of the
fore and middle legs with a few small but distinct spines ventrally, of the hind legs
with short, stout bristles, intermixed with which may be one or two spines. Lamellae
approximately one-fifth the length of the wing. Spermathecae two (in the single
preparation examined a third, very small, oval spermatheca was also present), highly
chitinised, oval, relatively large (64u x 36u), the commencement of the duct
chitinised for a very short distance.

Noé’s figure of the female palpi shows an exceptionally long, slender terminal
segment. This segment, if the drawing is accurate, is considerably longer than the
third (the ratio being 1:2: 1)—a condition which does not occur in any other species.
Unfortunately in the specimens at my disposal the palpi are absent or so damaged or
arranged that details cannot be observed.

Habitat. Italy: Roman Campagna; Sardinia, Cagliari. According to Weiss,
Bezzi believes that both L. ivritans and L. bezzii are widely distributed in Northern
Italy.

Leptoconops rhodesiensis, sp. nov.

Q.—Length of body, 2°55 mm.; length of wing, 1:2 mm.; length of antenna,
0'-4 mm.; width of head, 0°33 mm.

Head shining black, clothed with short, blackish hairs on the vertex and occiput ;
clypeus dark brown, with several (about twelve) dark-coloured hairs; eyes rather
widely separated, the space between them about one-third the greatest width of the
head. Proboscis dark brown or black. Palpi (fig. 2, c) dark brown, with dark hairs ;
third and fourth segments elongate, the third much swollen, with the orifice of the
sensory pit moderately large, subcircular, and almost centrally situated, the fourth
slightly longer than the third. Antennae (fig. 5, e) dark brown, with short paler brown
hairs and relatively stout blunt spines ; segments 4 to 13 transversely oval to sub-
spherical, the length being from 0:6 to 0°9 times the breadth ; fourteenth segment
ovate, as long as the two preceding segments together. Thorax shining black, clothed

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 15

with short black hairs ; scutellum and postscutellum similarly coloured, the scutellum
with three pairs of black bristles ; pleurae and pectus shining black. Wangs (fig. 7)
whitish, strongly iridescent ; first and third veins joining the anterior margin near
the proximal third, point of bifurcation of the fifth vein situated considerably beyond
the extremities of the costa and third veins. Halteres whitish, opalescent. Legs :
femora and tibiae dark brown, with pale hairs, metatarsi and second tarsal segments
without conspicuous spines, except distally, where differentiation (into spines or stout

Sw ey Sn,
= 5 NN
: : apenas SPS,

ATTN "

Fig. 7. Wing of Leptoconops rhodesiensis, sp.n., Q (x 75 circa). *

spine-like bristles) of the apical pair of bristles often takes place. Claws equal,
simple, each with a short bristle arising from the base. Abdomen : dorsum sepia-
coloured (shining when held in certain positions), the apical margins of the tergites
narrowly paler brown ; venter sepia-coloured medially, paler brown laterally ; both
surfaces with short dark hairs. Lamellae dusky white or light grey, 0°2 times as long
as the wing. Spermathecae strongly chitinised, apparently elongate-oval (collapsed
in the single specimen available) ; the commencement of the duct chitinised for
a very short distance.

Habitat. North-Western Rhodesia: Kafue Flats, 3,000 ft., 19.x.1913 (R. C.
Wood), ‘‘ Biting myself’’; 1 92 (type) in the collection of the Imperial Bureau of
Entomology.

This species should be recognised without difficulty by the wing venation, the
short extent of the anterior veins being a conspicuous character. From L. (Holoconops)
interruptus, End., the only other species yet described from South Africa, it may, of
course, be immediately separated by the structure of the antennae. The coloration
of the abdomen given above must be regarded with reserve, since the specimen was
caught when biting and probably contained undigested blood.

Leptoconops torrens, Twns.
Tersesthes torrens, Twns., Psyche, vi, pp. 369-371 (1893).

This species, the type of the genus Tersesthes, Twns., belongs to the group of species
in which the claws are simple. I am indebted to Professor L. O. Howard and
Dr. J. M. Aldrich for the opportunity of examining female specimens from the Organ
Mountains, New Mexico, and a microscopical preparation of a male from Las Vegas
Hot Springs, New Mexico. The females were collected by Townsend on horses, and
agree in all essentials with his description ; the males, however, cannot at present be
more than provisionally associated with this species, but in view of Weiss’s descrip-
tion of the male antennae of L. (Holoconops) kerteszi (M. laurae, Weiss), appear

16 HENRY F. CARTER.

to be referable to it rather than to the other American species—L. hertesz1 var.
americanus, NOV.

Q.—Length of body (one specimen), 2:2 mm. ;* length of wing, 1-1 mm. ; length
of antennae, 0°33 mm. ; width of head, 0°30 mm.

Head shining dark brown or blackish, the antennal depressions cinnamon-coloured,
clothed with short dark hairs on the vertex and occiput ; clypeus dark brown, with
twelve hairs—five on each side and two central; eyes separated by a Space equal to
about one-third the width of the head. Proboscis black. Palpi (fig. 2, 6) blackish
brown ; third segment greatly swollen, with the orifice of the sensory pit large, more
or less ovate, occupying the distal two-thirds of the inner side ; fourth segment sub-
cylindrical, distinctly (one-fifth) shorter than the third. Antennae (fig. 5, 2) short,
dark brown, with greyish hairs and moderately long, curved spines; fourth to
thirteenth segments transversely oval, from 0°6 to 0°8 as long as broad ; fourteenth
segment slightly more than twice as long as wide, equal in length to the three preceding
segments together. Thorax shining black or blackish brown, sparsely clothed with
short dark hairs ; scutellum with three pairs of black bristles, the central pair large.
Wings whitish, the anterior veins terminating in a brown stigma at some distance
before the middle, arranged as in fig. 8, e; fifth vein bifurcating below the ends of
the costal and third veins. Halteres white, the stems infuscated. Legs blackish
brown, the tarsi paler brown ; metatarsi and second tarsal segments without spines
except at the extremities. Claws simple, equal, each with a basal bristle. Addomen
brown, distinctly paler than the head and thorax, with short dark hairs. Lamellae
brown, 0°18 fhe length of the wing. Spermathecae two, highly chitinised, sub-
spherical (diameter 36) ; the commencement of the duct scarcely chitinised.

g—tLength of body (one specimen), 1‘°9 mm.; length of wing, 1-1 mm. ; length
of antennae, 0°71 mm.; width of head, 0°30 mm.

General coloration apparently (so far as can be judged from a balsam preparation)
dark brown or blackish, the tarsi paler. Head: frons bare, occiput and vertex with
scanty hairs ; clypeus with two pairs of short hairs, eyes widely separated, the space
between them being two-fifths the width of the head. Palpi: third and fourth
segments sub-equal, the third slender, the sensory organ situated in the distal half.
Antennae (fig. 4, h), very similar to those of the male L. bezztz (q.v.), but with the basal
segments of the flagellum less compressed, the fourteenth segment relatively shorter
(about three and one-half times as long as wide, and slightly more than one-third the
length of the last segment), and the fifteenth segment more strongly swollen distally ;
fourth to thirteenth segments varying from 0°6 to twice the width, fifteenth segment
nearly eight times as long as the greatest width (i.e., near the distal extremity),
fourteenth and fifteenth segments, taken together, about equal in length to the
preceding seven segments united. Thorax: scutellum with two pairs of bristles.
Legs slender, especially the middle and hind pairs; metatarsi of the four anterior
legs with a few small spines (usually one pair at the base and apex andone, unpaired,
near the middle), hind metatarsi and second tarsal segments with a pair of spines or
spine-like bristles at the apex. Claws of the fore and middle legs equal, one with a
long basal tooth (fig. 4, b), the other with a bristle ; of the hind legs equal and simple.
Hypopygium: Unfortunately the single preparation available is not in a sufficiently
good condition to allow a satisfactory interpretation of the detailed structure of
the intermediate appendages, but the claspers (fig. 4, a) are of peculiar form and
will probably provide specific characters.

* Townsend gives the length of the body (including the lamellae) as 1-6 mm. to 2-2 mm,
according as the abdomen is empty or distended with blood.

+ The characters afforded by the wings cannot be determined in microscopical preparations
with any degree of accuracy, unless the specimen is stained. In addition, the wings in the
specimen described were considerably twisted.

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. NL

Habitat. U.S.A.—New Mexico: Continental Divide, 7,000 ft., 21st June,
(C. H. T. Townsend—type series), Organ Mountains, 5,700 ft., 29th Aug. (C. H. T.
Townsend— 2 described above) ; Las Vegas Hot Springs (H. S. Barber— 3 described
above).

Dr. Aldrich informs me that, besides the localities mentioned above, the Leptoconops
material in the United States National Museum includes females from Arizona,
Colorado, Florida, Texas, Utah and Cuba, and males from Arizona. The specimens
from Utah are L. kerteszt var. americanus, nov., but the others have not yet been
definitely determined, and are provisionally referred to L. torrens.

Pratt (1907) mentions several of the foregoing localities, including Utah, in
connection with L. torrens.

Leptoconops bezzii, Noé.

Mycterotypus bezzi1, Noé, Atti R. Accad. Lincei, Ser. 5, Rendiconti xiv, p. 114
(1905) ; Arch. Zool. Napoli, iii, p. 187 (1907).

Leptoconops hyalinipennis, Kieft., Ann. Mus. Nat. Hung. xvi, p. 33 (1918).

The following description of L. bezzit (2) is drawn up from specimens collected in
Central Italy and sent me by Professor M. Bezzi; to this gentleman I am also
indebted for the loan of microscopical preparations of the palpi and antennae of
the type male.

?.—Length of body (two specimens), 2-1 mm. ; length of wing, 1-2 mm. ; length
of antenna, 0-56 ; width of head, 0-45 mm.

Head black, clothed on the vertex and occiput with short dark hairs; clypeus
black or blackish brown, with two pairs of short dark hairs ; eyes moderately widely
separated, the space between them about one-quarter the width of the head; frons
with a pair of short hairs near the lower margins of the eyes. Proboscis dark brown,
Palpi dark brown, with dark hairs ; third segment strongly incrassate, the orifice of the
sensory pit very large, narrowly oval, extending almost the entire length of the inner
side; fourth segment subcylindrical, slightly shorter than the third. Antennae
dark brown, with short dark hairs and clear spines, which are somewhat strongly
curved on the distal segments ; fourth to thirteenth segments transversely oval to
subspherical, the length from 0-8 to 1-0 the breadth ; fourteenth segment about
two and one-quarter times as long as broad, equal in length to the preceding
two and one-third segments together. Thorax black, sparsely clothed with
short black hairs; scutellum black, with three pairs of dark bristles (one of the
small lateral bristles sometimes wanting) ; postscutellum, pleurae and pectus black.
Wings white, iridescent, the anterior veins arranged as in fig. 8, f; fifth vein bifurcating
slightly beyond the end of the costa. Halteres white. Legs dark brown or brownish
black, the metatarsus and second tarsal segment paler, yellowish brown, each with a
pair of apical spines. Claws (fig. 4, d) equal, each with a stout basal tooth. Abdomen
dark brown, clothed with short black hairs. Lamellae yellowish brown, 0-2 the
length of the wing. Spermathecae two, heavily chitinised, oval (70m by 54”); the
commencement of the duct only chitinised.

The male obtained and associated with this species by Noé differed from the
female chiefly in regard to the palpi, antennae, claws and wings. The antenna
consists of fifteen segments, the third (i.e. the first segment of the flagellum) being
produced basally into a relatively long stalk, and causing the flagellum to be more
distinctly separated from the basal segments than in the female: the fourth to
thirteenth segments become progressively longer and narrower, the most proximal
segment being very short and broad (0-4 to 0-7 the length), with chitinous thickenings
on the distal margins, the thirteenth subspherical basally but produced anteriorly

(2416) B

18 HENRY F. CARTER.

(the greatest width being 1-8 the length) ; the fourteenth and fifteenth segments
greatly elongate, the former about half the length of the latter, and when united
almost equal in length to the remaining segments of the flagellum taken together.
The palpi are longer and more slender than in the female, the third segment elongate,
scarcely swollen, with the proximal two-thirds of the inner side excavated, the fourth
segment somewhat swollen distally, approximately four-fifths the length of the third.
The wings are more delicate, and (from Noé’s figure, not his interpretation) the first
and third veins appear to be fused basally, and to enclose a large interspace distally ;
the separate distal portion of the first vein is very short and directed abruptly upwards
towards the costa, the separate portion of the third vein long, extending at first
almost parallel with the costa, then curving gradually upwards to meet it a short
distance beyond the middle of the anterior border and above the bifurcation of the
fifth vein. The claws, according to Noé, are dissimilar, those on the anterior legs
being provided with a long basal tooth, those on the hind legs simple, with a short
basal bristle.

Kkieffer’s description of L. hyalinipennis agrees so closely with Noé’s description
of L. bezzit, and with the specimens at my disposal, that I have no hesitation in placing
it as a synonym of the latter species.

Habitat. L. bezzi1 is now known to occur in the Roman Campagna, Central Italy,
and (as L. hyalintpennis) in Tunis (Djebel Djeloud ; Korbons ; Aouina, Lac Bahira ;
Tunis, Pare Belvedere).

Leptoconops flavivertris, Kieff.
Leptoconops flaviventris, Kieff., Ann. Mus. Nat. Hung. xvi, pp. 34 and 85 (1918).
Kieffer’s description of the female of this species is as follows :—

“©—Semblable a L. hyalinipennis, sauf les caractéres suivants: Bouche plus
longue que la hauteur de la téte, dirigée en arriére. Palpes de 3 articles, dont le ler
est mince et un peu plus long que gros, seulement un article aprés laflexion, comme
chez hyalinipennis. Antennes a articles 3-12* trés transversaux, soies sensorielles
plus courtes que les poils des verticilles, 13e en ovoide allongé, sans verticille,
au moins aussi long que les précédents réunis. Mesonotum luisant. Ailes blanches,
nervures tres pales. Tarses blanchatres, articulations sombres. Abdomen jaune
soufre. L. 1-3 mm.

Djebel Tunisie. Djeloud (59).”’

On a later page Kieffer recorded additional specimens, including a male from
Asia Minor—“ Kyaldja-Su, viii (Naday), 1g; Emirley; Suleyman, H. Yayla;
Kkarapunat, 7 9 (Naday).’’ The male, which he doubtfully associated with this species,
is shining black, with brown legs and a pale brown abdomen, but from the description
given it is difficult to select any very definite characters by means of which it may be
distinguished from the male of L. bezzii. Kieffer, however, states that the proboscis
is very long and slender—much longer than the height of the head—and that the
metatarsi are devoid of spines. Segments four to thirteen of the antenna appear to be
similar in form to those of the Italian species, but the distal portions of these organs
were evidently damaged, and Kieffer was unable to determine whether fourteen or
fifteen segments were present. The claws are merely described as long (more than
half the length of the fifth tarsal segment), but apparently the association with L.
jlaviventris would imply that on some of the legs, at least, they are also toothed.

* In this and subsequent direct quotations the number of the antennal segment given by the
author cited must, for correctness and uniformity, be increased by one ; but in all the descriptive
extracts given this change has already been made.

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 19

Of the species of Leptoconops occurring in the Mediterranean littoral L. flaviventris
apparently most closely resembles L. ivritans in general facies. In both species the
abdomen is normally pale in colour—whitish or yellowish—but they should be easily
separated by the structure of the claws.

HFC

Fig. 8. Basal portions of wings of females of: (a) L. stygius, Sk. ; (b) L. longicornis, sp. n. ;
(c) LE. grandis, sp. n.; (d) L. stamensis, sp. n.; (e) L. torrens, Twns.; (f) L. bezzi, Noé; (g)

L. braziliensis, Lutz; (h) L. herteszi, Wieff.; (k) L. kerteszi var. americanus, n. ; (1) A. spinosifrons,
Spyms (Orca, <x o0/-circa) the rest: x 7).

Leptoconops indicus, Kieff.
Schizoconops indicus, Kieff., Ann. Mus. Nat. Hung. xvi, p. 135 (1918).

This species was made the type of the genus Schizoconops by Kieffer on account of
its toothed or bifid ungues. This character alone, however, cannot be considered of
sufficient importance to warrant the creation of a new genus; and even were it of
subgeneric value the name Schizoconops would sink under Mycterotypus (see p. 3).

The chief characters, taken from Kieffer’s description, are as follows :—

9.—Shining black. Antennae brown, the fourth to the thirteenth segments
transverse, at least twice as broad as long; fourteenth segment conical, equal in
length to the four preceding segments together. Wings white, with pale veins ;
first and third veins not reaching the middle of the anterior border, fifth vein
bifurcating much beyond the extremity of the third vein. Halteres white. Legs pale
(2416) B2

20 HENRY F. CARTER.

brown, tarsi whitish, metatarsi without spines ; anterior femora slightly thickened ;
claws bifid, equal, the branch shorter. Abdomen red ; lamellae long and whitish.

Length 1-5 mm.
Habitat. Bengal: Champaran, Bettiah, iti, 1908.

This species should be easily recognised by the structure of the claws, the absence
of spines on the metatarsi, and the relative positions of the extremity of the third vein
and bifurcation of the fifth vein ; that the latter should take place considerably beyond
(‘‘trés distale ’) the junction of the third vein with the costa is unusual.

Leptoconops siamensis, sp.nov.

9.—Length of body, 3-5 mm.; length of wing, 1-8 mm.; length of antenna,
0-6 mm. ; width of head, 0:43 mm.

Head dull brown, the antennal depression creamy-white, clothed with dark brown
hairs on the occiput ; clypeus rather paler brown, with four dark brown hairs on each
side of the middle line ; eyes relatively not very widely separated, the space between
them almost one-fifth the greatest width of the head and devoid of hairs. Proboscis
pale brown. Palpi (fig. 2, /) pale brown, with brown hairs ; third and fourth segments
elongate, subequal ; the third strongly incrassate, with a deep sensory pore, the orifice
of which is very large and occupies almost the entire length of the inner side.
Antennae moderately long, yellowish brown, with short pale brown hairs and slightly
curved, pointed, transparent spines; segments 4 to 13 subspherical to oval, the
length varying from 0-8 to 1-3 times the breadth; terminal segment (fig. 5, a)
elongate, equal in length to the preceding two and one-third segments together.
Thorax : disc dark umber-brown, pollinose, clothed with short brown hairs; pro-
thoracic lobes and humeral callus yellowish brown ; scutellum and postscutellum rather
darker than the disc, the former with three pairs of strong median bristles and two
pairs of small lateral hairs ; pleurae and pectus dark umber-brown. Wangs whitish,
venation normal, the first and third veins (fig. 8, d) fused distally, not forming an
interspace. Legs entirely brownish yellow, bearing pale brown hairs and, on some of
the tarsal segments, stout blackish spines; fore femora and tibiae slightly swollen
and somewhat shortened, tibia of all the legs with an apical spur; metatarsi each
with two sub-ventral or ventro-lateral rows of strong spines,* the second and third
tarsal segments of the fore and middle legs each with two apical spines, of hind legs
wanting. Claws (fig. 4, f) of fore and middle legs equal, each with a large (at least
half the length of the claw) strong tooth arising from the base. Abdomen waxen
creamy white above and below, clothed with short hairs. Lamellae brownish yellow,
approximately 0-25 times the length of the wing. Spermathecae two, rather narrowly

oval (654 x 38y), highly chitinised ; commencement of the duct chitinised for a very
short distance.

Habitat. Siam: Patani Cape (H. C. Robinson & N. Annandale). One female
(type) in the British Museum Collection.

This species is not closely allied to any of the known members of Leptoconops
(sens. lat.) ; it agrees with L. indicus in regard to the structure of the claws, but
may readily be separated therefrom by its relatively.large size and the powerful
spinose armature of its metatarsi. In the latter character and the reduction in length
of the fore legs it suggests Acanthoconops (q.v.).

* The development of these spines has not proceeded uniformly, and although normally paired,
the spine on one side is often much smaller than that on the other and may be represented by a
strong bristle. The number of spines present on the same segments of corresponding legs there~-
fore varies, and on the metatarsi of the single specimen available was—fore legs 18 and 15, middle
legs 14 and 15, hind legs 9 and ? (the metatarsus of the corresponding leg missing).

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. PA

Subgenus Holoconops, Kieff.
Holoconops, Kieff., Ann. Mus. Nat. Hung. xvi, p. 135 (1918).

This subgenus comprises those species of Leptoconops (sens. lat.) in which the
antennae of the female are composed of thirteen segments ; in all the known species
the claws in this sex are simple and equal. The group was given generic rank by Kieffer,
who based its separation upon the structure of the female antennae and claws. Sucha
combination of characters, however, cannot be maintained, since the structure of the
claws is in no way pectiiar, and this author’s restriction of the type mentioned above
to Holoconops is evidently due to a misconception. Kieffer designated L. kerteszi,
Kieff., as his genotype, and, in a footnote, associated it with L. flaviventris, Wieff.,
L. hyalinipennis, Kieff. (synonymous with L. bezzit, Noé), and L. lacteipennis, Kieff. ;
but the inclusion in this group of the second and third-named species is a palpable
oversight, as in his descriptions of them on preceding pages of the same article the
antennae are definitely stated to possess the full complement of segments (fourteen).

Three species of this subgenus are here recognised, but one (L. interruptus, End.)
is insufficiently described, and may subsequently prove identical with one of the
others. They are widely distributed, and have been recorded from Northern Africa,
Asia Minor, South West Africa and the United States of America.

Leptoconops kerteszi, Kieff.
Leptoconops kerteszi, Kieff., Ann. Mus. Nat. Hung. vi, p. 576 (1908).
Mycterotypus laurae, Weiss, Arch. Inst. Pasteur de Tunis, pp. 25-32 (1912).

Mycterotypus laurae var. peneti, Langeron, Arch. de Parasit. xvi, pp. 282-301
(1913).

The synonymy given above is based upon examinations of specimens sent me as
L. kerteszi by Mr. F. C. Willcocks from Egypt, and of females of L. lawrae sent me by
M. A. Weiss from Tunis. L. kerteszi was described by Kieffer from specimens collected
at Cairo, and the material received from Mr. Willcocks agrees in detail with this
author’s descriptions. Furthermore Kieffer, who is evidently unacquainted with
L. laurae, has recently (1918) recorded L. kerteszi from Tunis.

The following description is compiled from the Egyptian and Tunisian specimens
referred to above.

°.—Length of body (six specimens), 1-8-2-1 mm.; length of wing, 1-1-1-3 mm. ;
length of antenna, 0-33-0-42 mm. ; width of head, 0:31-0:34 mm.

Head shining black, the antennal depressions pale buff, sparsely clothed on the
vertex and occiput with short black hairs ; clypeus shining black, with three pairs of
dark hairs ; eyes relatively widely separated, the space between them being approxi-
mately two-fifths the width of the head. Proboscis dark brown or black. Palpi
(cf. fig. 2, d) dark brown, with brown hairs ; third segment very strongly swollen,
with a large oval pore situated near the middle, fourth segment slightly inflated
distally, scarcely shorter than the third. Antennae (cf. fig. 5, g) dark brown, with
rather long (about twice the length of the segment) pale brown hairs ; fourth to twelfth
segments transversely oval to spherical, from 0-8 to 1-0 as long as broad ; thirteenth
segment equal in length to the preceding three and one-third to four segments together.
Thorax entirely shining black, with short black hairs ; scutellum with two pairs of
bristles. Wings white, iridescent ; venation normal, the anterior veins not quite
reaching the middle of the anterior border and arranged as in fig. 8, ; fifth vein
bifurcating slightly before the extremities of the costa and third veins. Halteres
whitish. Legs dark brown, clothed with dark hairs ; metatarsi of the hind legs paler
brown, with short brownish yellow hairs ; fore and middle metatarsi with a basal and
apical pair of slender, pointed spines, and a few (one to four) central, usually unpaired,

22, HENRY F. CARTER.

spines ; hind metatarsi and second segments of all the legs with a pair of similar spines
at the extremities. Claws (cf. fig. 4, c) simple and equal, each with a short bristle
arising from the base. Abdomen dark brown, the margins of some of the tergites
narrowly grey, sparsely clothed with short brown hairs. Lamellae pale brown, 0-25
the length of the wing. Spermathecae two (a third very small, narrowly oval one is
often present), heavily chitinised, obovate, (47 36) ; a minute portion only of the
duct chitinised.

The male was described by Weiss, who stated that it was uniformly darker in
colour than the female, and gave the following measurements :—Length of body,
1-5mm.; length of antennae (barely), 0-5 mm. Judging by this author’s description
and figures, it presents certain striking morphological differences from those males
which have been associated with species of Leptoconops (sens. str.). The eyes are said
to meet at a point above ; the palpal segments are depicted as subequal in length and
about three times as long as wide ; the fourth antennal segment almost twice as long
as the fifth, the fifth to twelfth short and broad, subequal, the last three segments
elongate, but the thirteenth almost equal in length to the fourteenth and fifteenth
taken together ; and the wings with reduced venation—the fourth vein being absent.

Habitat. Egypt: Cairo, Behera, Wadi Natroun, Sakkara (Willcocks). Tunis:
Tabeditt (Weiss).

Willcocks (1917) states that the species was first sent from Behera in March 1907,
and that it is common at certain seasons in Wadi Natroun, and in the autumn, when
the Nile is in flood, from Mena House to Sakkara.

Leptoconops kerteszi var. peneti, Langeron.

°.—Length of body, 1-5-1-8 mm. ; length of wing, 1:06 mm. ; length of antenna
(from figure), 0-45 mm.

According to Langeron this variety differs from the specimens (type series) of
M. laurae in the Paris Museum principally in being larger, darker, and possessing more
numerous and stronger bristles and spines. In particular, he compares the develop-
ment and exact arrangement of the metatarsal spines in the two forms. He also
maintains that biological differences exist: the variety appears to persist Jater in the
year, occurs in a more southerly region, and at a much lower altitude than is recorded
by Weiss for the type form. MW. laurae came from a mountainous region (altitude
500 metres), while the var. penet? was found in the desert (mean altitude 30 metres),
in the marshes lying between the oases of E] Hamma and the Shott Gharsa.

In view of the individual variation in regard to size, and number and arrangement
of the metatarsal spines observed in a series of specimens of L. kerteszi, the distin-
guishing points cited by Langeron can scarcely be granted even varietal value ; but
in this author’s excellent and detailed description of his specimens, mention is made
of a character which is of much greater importance, and which raises doubt regarding
its specific identity. The ungual formula is stated to be 0-1—0-1—0-1, and the
external claw of each leg is said to bear a small basal tooth. Such a condition occurs
in no other species of Leptoconops, and therefore, if Langeron’s interpretation be correct,
the form should be easily recognised and would deserve specific rank. It should be
noted, however, that in some species the base of the claw is relatively broad and
projects slightly ventrally, so that, in certain positions, a minute basal tooth appears
to be present.

Leptoconops kerteszi var. americanus, nov.

9.—Length of body (three specimens), 1-75 mm.; length of wing, 1-1 mm. ;
length of antenna, 0-32 mm. ; width of head, 0-30 mm.

In spite of their widely distant places of origin I have been unable to find any
satisfactory characters for separating specimens from Utah (received through the

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 23

kindness of Professor L. O. Howard and Dr. J. M. Aldrich) from typical examples of
L. kerteszi. Indeed, allowing for minute differences which come within the range of
individual variation, the only distinguishing details appear to be the formation of an
interspace by the first and third veins (fig. 8, &) and the slightly different form of the
spermathecae. These are almost spherical in the Utah specimens, whereas they are
obovate in the Mediterranean examples. Males of both forms, when forthcoming and
compared, may possibly reveal distinctive antennal or genital characters, but until
such time the American form can, at the most, be accorded varietal rank.

Habitat. United States of America: Utah, Salt Lake, June, “ biting devilishly ”
(H. S. Barber). Three females (cotypes) in the collections of the United States
National Museum and Liverpool School of Tropical Medicine.

Leptoconops lacteipennis, Kicff.
Leptoconops lactetpennis, Kieff., Ann. Mus. Nat. Hung. xvi, p. 32 (1918).

This species, described from a female captured in Tunis, apparently difters from
L. kerteszi chiefly in the metatarsi being devoid of spines, the segments of the flagellum
more uniformly transversely oval, and the fifth tarsal segment Shorter than the fourth.
The last character should enable the species to be identified without difficulty, since,
so far as I am aware, it is unique in this respect.

The more important of the specific characters given by Kieffer are as follows :—

9.—Black. Fourth to twelfth antennal segments transverse (from the figure
accompanying the description, the length varies from 0-6 to 0-7 times the breadth),
thirteenth segment slightly longer than the three preceding segments together.
Mesonotum shining. Halteres white. Wings milky white, extending almost to the
tip of the abdomen ; third vein reaching almost to the middle of the wing, fifth vein
bifurcating slightly before the extremity of the third vein. Legs black, without
spines, tarsi brown ; tarsal segments 1 to 4 gradually decreasing in length, the fifth
segment shorter than the fourth. Claws simple, equal, each with a bristle arising
from the base. Abdomen brownish black. Length, 2 mm.

Leptoconops interruptus, End.
Mycterotypus interruptus, End., Denks. Med. Ges. Jena, i, pp. 133-162 (1908).

The description of this species given by Enderlein relates largely to characters of a
general nature, and details which, in the hght of our present knowledge of the group,
must be considered of value in separating such closely allied forms receive little
attention. Nevertheless, with the help of the figures accompanying the description,
it would appear that L. interruptus is very close to, if not identical with, L. kerteszt.
Weiss (1912) observed the close relationship existing between these species, and
especially noted the affinities existing in regard to the spinose armature of the
metatarsi and the structure of the antennae. Since, however, the exact arrangement
of the metatarsal spines is not of specific importance, it seems that the only
tangible differential character is afforded by the antennae. [nderlein states that
the third to twelfth segments are spherical, and that the last segment is twice as
long as broad; this would indicate, also, that the terminal segment was approximately
equal in length to the two preceding segments together. In L. kerteszi the last
segment is at least three times as long as broad, and is equal in length to the three to
four preceding segments.

A re-examination of the type or the examination of further material from South-
West Africa (Enderlein’s example came from Rooibank, hinterland of Walfish Bay) is
necessary, before a decision regarding the validity of one or both of these species can
be made.

24 HENRY F. CARTER.

Genus Acanthoconops, nov.*

Frons clothed with bristles or spines. Antennae in the female pilose, composed
of fourteen segments, the fourth to thirteenth short and broad, subspherical, the
fourteenth elongate, subconical. Eyes, palpi, proboscis and wings as in Leptoconops,
Sk. (sews. lat.). Fore legs relatively short, the fore and hind femora and tibiae
strongly incrassate, the middle femora moderately incrassate; claws equal, each
with a tooth arising from the base ; empodium bristle-like. Ovipositor somewhat
triangular, very short, considerably broader than long.

Genotype: A. spinosifrons, sp. nov.

This genus is very closely allied to Leptoconops (sens. lat.), but may readily be
distinguished by the remarkably short ovipositor (fig. 9), and by the vestiture of the
head. In Leptoconops (sens. lat.) the whole of the frons (i.e., the wide area extending
from the vertex to the clypeus) is bare, or, at most, possesses a single pair of bristles
between the eyes, while in Acanthoconops it bears numerous spines or bristles (Cie
fig. 1,a@ and 6). Further, in the two species of Acanthoconops at present known the
fore legs are noticeably shortened, and the metatarsi of all the legs armed with
formidable spines. The ratio of the combined lengths of the femora and tibiae of the
fore legs to those of the hind legs is 1: 1-5, or but slightly less ; in Leptoconops this
ratio is rarely more than | : 1-2, and not infrequently is 1:1. In this connection it
may be of interest to note that, among the members of the latter genus, the ratio
given for A canthoconops is (so far as can be determined from the material available)
most nearly approached in the case of L. siamensis, sp. n. In this species, which also
possesses powerful spines on the metatarsi, the ratio is almost 1 : 1-4.

Fig. 9. Extremity of abdomen of Acanthoconops spinosifrons, sp. n., 9, showing lamellae
(2); side view (x 220 circa).
Acanthoconops spinosifrons, sp. nov.
?.—Length of body (two specimens mounted in Canada balsam), 2:3 mm. ;
length of wing, 1-2 mm. ; length of antenna, 0-35 mm. ; width of head, 0-32 mm.

Head dull black, with short, stout, pointed, backwardly directed, tuberculate
spines scattered over the occiput and frons, on the lower portion of which they are
more numerous ; clypeus shining dark brown, with a group of spines centrally and a

* axavOa spine, and x@vow gnat.

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 25

few (three pairs) relatively short black hairs ; eyes widely separated above and below,
the space between them at the vertex being rather more than one-third the greatest
width of the head. Proboscis dark brown. Palpi (fig. 2, e) dark brown, the apical
segment yellowish brown, clothed with short dark hairs; apparently composed of
three segments, the two small basal segments being almost fused together ; orifice
of the sensory pit in the third segment small, subcircular, situated in the
proximal third. Antennae (fig. 5, f) relatively short, dark brown, with short glistening
hairs and short, straight or slightly curved spines; segments 4 to 13 transversely
oval to subspherical, the length varying from 0-6 to 0-8 of the breadth, the fourteenth

fae

Fig. 10. Acanthoconops spinosifrons, sp.n., 2: a, front leg (x 85); }, first and second tarsal
segments of front leg (x 295); c, hind leg (x 85); d, first and second tarsal segments of hind
leg (x 295) ; e, fifth tarsal segment of hind leg (x 550).

segment equal in length to the preceding two and one-half segments together. Thorax
and scutellum shining black, the dorsum clothed with short brown hairs ; scutellum
bearing two pairs of large, and one or two pairs of small bristles ; pleurae and pectus
shining black. Wings glassy, strongly iridescent ; the anterior veins as shown in
fig. 8, /, the fifth vein bifurcating almost immediately below the extremity of the
third vein. Halteres with white knobs and infuscated stems. Legs (fig. 10) pale
brown, the distal two-thirds or three-fourths of the femora, and the distal third of the
fore and hind tibiae, black, clothed with brown hairs; tibiae each with a strong
apical spur, fore tibiae incrassate, hind tibiae with five strong, blunt, black spines on

26 HENRY F. CARTER.

the outer side at the apex, and with three of the bristles of the posterior transverse
distal row (situated on the inner aspect) replaced by similar but smaller spines ;
metatarsi, particularly those of the hind legs, short, fore and hind metatarsi incrassate ;
fore and hind tarsi with stout, blunt black spines arranged as shown in fig. 10,
middle tarsi with six or seven pairs of similar spines on the first segment and the
apical bristles of the second and third segments differentiated—spine-like. Claws
(fig. 10, e) equal, the basal tooth stout. Abdomen translucent creamy white, but
appearing in engorged or partly fed specimens dark brown or cream-coloured, with
dark central bands on the proximal segments; clothed with pale hairs. Lamellae
creamy white, clothed with pale hairs. Spermathecae two in number, heavily
chitinised, oval (53437) ; the commencement of the duct chitinised for a
relatively long distance (18y).

Habitat. Zanzibar (Dr. W. M. Aders), seven females (including three cotypes).
The labels attached to the specimens bear the following data: on buffalo ; Pigaduri,
Zanzibar, 13.iv.19. In the collection of the Imperial Bureau of Entomology.

Acanthoconops albiventris, de Meijere.
Leptoconops albiventris, de Meijere, Tijds. voor Ent. lviii, p. 98 (1915).
Leptoconops spinosipes, Kieff., Ann. Mus. Nat. Hung. xv, p. 190 (1917).

?.—Length of body (two specimens), 1-8 mm.; length of wing, 1-0 mm. ;
length of antenna, 0-3 mm.; width of head, 0-32 mm.

Through the courtesy of Professor de Meijere, I have been able to examine females.
of this species, which was described by him from specimens collected in New Guinea.
This author noticed and commented upon the unusual form of the lamellae, but
retained the species in Leptoconops, and apparently did not observe closely the
arrangement of the hairs on the head ; this he stated was ‘“ kaum_ behaart.”’
A. albiventris is closely allied to the preceding species, but is smaller, and possesses.
somewhat less powerful, though similarly arranged, spines on the legs. Morpho-
logically, it may readily be separated from A. spinosifrons by (1) the frons bearing
numerous short hairs or bristles instead of spines ; (2) the terminal segment of the
antennae being relatively longer (equal in length to the preceding three and one-
third segments instead of the preceding two and one-half) ; (3) the bristles forming
the posterior transverse distal row on the hind tibiae all being normal, none replaced
by spines ; (4) the hind metatarsus being relatively longer (about one-half the length
of the tibiae, whereas in A. spinosifrons it is about one-third the length) ; and (5) the
tooth of the claws being distinctly smaller.

The synonymy given above seems extremely probable from a comparison of the
descriptions, and in view of the fact that Kieffer’s specimens also came from New
Guinea (Tamara, Berlinhafen). Discrepancies in the descriptions are slight, and
Kieffer’s statement that the abdomen is red, sometimes white, with brownish
markings, is of little consequence, as it suggests that some (possibly most) of his
examples were wholly or partly engorged with blood.

This species appears to be a vicious biter and, at times, a serious pest in parts of
New Guinea; K. Gjellemp, the collector of de Meijere’s material, attached the
following information to the specimens— An der Mundung des Sermowai-Flusses

in sehr grosser Anzahl vorhanden und durch ihr Stechen eine grosse Plage bildend,
16 Mai 1911.”

The distinguishing characters of the known species (females) of Leptoconops (sens.
lat.) and Acanthoconops are summarised in the following table.

(1) Lamellae elongate ; frons bare or with a single pair of hairs between the
eyes (Leptoconops, sens. lat.) i ae ; ; : bi

Lamellae very short ; frons with numerous spines or hairs (Acanthoconops) 17

A REVISION OF THE GENUS LEPTOCONOPS, SKUSE. 2T

Antennae composed of fourteen segments (Leptoconops, sens. str.) .. stages

Antennae composed of thirteen segments (Holoconops, subgen.) .. Heo 3
Claws simple oh: we © if ae ie. ie a ey
Claws toothed .. a - Af 6 a a ne eee
Metatarsi with distinct spines .. S 4 2 oie Ae a ae ee
Metatarsi without spines (excluding the pair at the apex) .. ae a 6
Abdomen whitish or yellow ; lamellae approximately one-fifth the length

of the wing .. me of ae es oie ivritans, Noé (p. 14).

Abdomen dark brown; lamellae approximately one-third the length of
the wing Me at Be oe as .. braziliensis, Lutz (p. 13).

Fifth vein bifureating considerably beyond extremity of costa : :
rhodesiensis, sp. n. (p. 14).

Fifth vein bifurcating before or bélow extremity of costa .. au ; a

Third palpal segment greatly swollen; antennal segments 4-13 copoly
broader than long (American species) Ae .. torrens; Twns. (p. 15).

Third palpal segment elongate, slightly swollen ; antennal segments 4-13
subspherical or longer than broad (Australian SPeGles) "lie a tie = 20

Smaller species (wing length 1.8 mm.) ; last antennal segment less than
twice as broad as long 7 23 q; .. Stygius, Skuse (p. 10).

Larger species (wing length 2:0 mm.) ; last antennal segment more than
three times as broad as long 2; Le as - : Be ee Me.

Antennae very long, last segment at least six times as long as Bree
longicornis, Spi pay

Antennae shorter, last segment at most three and one-half times as broad

as long.. od a 6 st: <4 oi prandis, sp. mn. (pa b2)e
Metatarsi with very large spines .. Slamensis, sp. n. (p. 20).
Metatarsi without spines (excluding the pair ‘at the apex) he st tale
Antennal segments 4-13 transversely oval to spherical, the last segment

slightly longer than the two preceding together .. bezz11, Noé (p. 17).
Antennal segments 4-13 all broader than long, the last segment as long as

the three or four preceding together ae = : oe sir OR
Abdomen yellow ; last antennal segment as long as the ae preceding

together a flaviventris, Kieff. (p. 18).

Abdomen red ;_ last cell Scrat as Sian as the four preceding
indicus, WKieff. (p..19).
Metatarsi without spines ; fifth tarsal segment shorter than fourth 4.1
lactetpennis, Kieff. (p. 23).
Metatarsi with distinct spines ; fifth tarsal segment longer than fourth .. 14
Last antennal segment twice as long as broad, segments 4-12 spherical ..
interruptus, End. (p. 23).
Last antennal segment three to four times as mi as broad, segments 4-12
transversely oval to spherical aye o ea eo)
Claws equal, one simple, the other with a miei BaeAll tooth oe
kerteszi var. penett, Langeron (p. 22).
Claws equal and simple 1

Spermathecae obovate, narrow Aenale (Medien ane ceeon)
kerteszt, Kieff. (p.

21
Spermathecae subspherical (America) kerteszi var. americanus, n. (p. 22).

28 HENRY F. CARTER.

(17) Frons spiny (Tropical Africa) .. bis Sate spinosifrons, sp.n. (p. 24).
Frons hairy (New Guinea) - se sit albiventris, Meij. (p. 26).

. References.
CASTELLANI, A. & CHALMERS, A. J. (1913). A Manual of Tropical Medicine.
London, 2nd ed., p. 678.
si . (1919). -. Wid. 3rdied.> pr S03:

CHATTON, E. & Branc, G. (1917). Notes et réflexions sur les Toxoplasmes et la
Toxoplasmose du Gondi (Toxoplasma gundit, Nicolle et Marceaux, 1909).—Arch.
Inst. Past. Tunis, x., pp. 1-40.

GrassI, B. (1901). Die Malaria: Studien eines Zoologen. Jena, pp. 118-122.

JOHANNSEN, O. A. (1905). Aquatic Nematocerous Diptera, II—N.Y. Sta. Mus.
Bull. No. 86, pp. 94-96.

KIEFFER, J. J. (1906). Chironomidae. Genera Insectorum. Wytsman. 42efasc.

Diptera, pp. 44-48.

(1908). Description d'une espéce nouvelle de Chironomides d’ Egypte.
—Ann. Mus. Nat. Hung. vi.

(1917). Chironomidae d’Australie.— Ibid. xv, pp. 189-191.

(1918). Chironomides d’Afrique et d’Asie.— Jbid. xvi, pp. 31-136.

LANGERON, M. (1913.) Mvycterotypus laurae; description d’une variété nouvelle
(M. laurae var. peneti).—Arch. Parasit. xvi, pp. 282-304.

Lutz, A. (1912). Contribuicgdéo para o estudo das Ceratopogoninas haematofagas do

Brazil.—Mem. Inst. Osw. Cruz, iv, p. 24.
(1913). Do.— Ibid, v, pp. 45-72.

Mattiocu, J. R. (1915). The Chironomidae, or Midges, of [linois, with particular
reference to the species occurring in the Illinois River.—Bull. Ill. Sta. Lab. Nat.
Est x, Ant, 6,7. 401

MEIJERE, J. C. H. de (1915.) Diptera aus Nord-Neu Guinea.—Tijds. voor Ent.
lviii, pp. 98-101.

Mik (1894). Wiener Ent. Zeit. xiii, p. 164.

Nok, G. (1905). Un nuovo genere appartenente alla famiglia Chironomidae.—

Atti. della Reale Accad. dei Lincei, Ser. 5, Rendiconti, xiv, pp. 114-120.
(1907). Due nuove specie di Ditteri appartenenti ad un genere nuovo.—
Arch. Zool. Napoli, iii, pp. 101-164.

PratTT, F.C. (1907). Notes on “ Punkies.’’—U.S. Dept. Agric. Bur. Ent. Bull. No. 64,
part 3, pp. 23-28.

SAMBON, L. W. (1913). The Causation of Pellagra : a Contribution to the Discussion
on Dr. Sandwith’s Paper.—Trans. Soc. Trop. Med. & Hyg. vi, pp. 231-241.

Skuse, A. A. (1890). Diptera of Australia. Part VI. The Chironomidae.—Proc.
Linn. Soc. N.S.W. (2nd Ser.), iv, p. 288.

Townsenb, C. H. T. (1893). An interesting Blood-sucking Gnat of the family
Chironomidae.—Psyche, vi, p. 369-371.

Wess, A. (1912). Mycterotypus laurae, n. sp., Chironomide nouveau du Sud Tunisien.
—Arch. Inst. Past. Tunis, pp. 25-32.

Wittcocks, F. C. (1918). Notes on some Insects found in Egypt of Medical and
Veterinary Interest.—Bull. Soc. Ent. d’Egypte, 3rd fasc., July-Sept., p. 84.

’

20
MOSQUITO BREEDING IN SALINE WATERS.

By ANDREW BALFouR, C.B., C.M.G., M.D.

Director-in-Chief, Wellcome Bureau of Scientific Research.

In the “ Bulletin of Entomological Research ” for December 1920 Dr. J. M. Dalziel
deals in a most interesting and instructive way with the various, and sometimes curious
breeding-places of mosquitos in Lagos. He mentions the occurrence of mosquito
larvae in saline waters, and in a foot-note refers to a number of salt-water species.

The subject of mosquitos breeding in salt and brackish waters possesses not only
considerable scientific interest, but is of practical importance in anti-malarial work,
for the suggestion has frequently been made to abolish ordinary mosquito breeding-
places by the introduction of salt or of sea-water, and in some instances this procedure
has actually been carried into effect. As will be seen, in the case of certain species of
mosquitos it is useless or worse than useless.

Moreover, there is the question of the efficiency of larvicides in saline waters, a
matter to which attention has been directed, but on which, so far as I know, our
information is still defective.

It may therefore be of interest to refer a little more fully to the subject than
Dr. Dalziel has done, although at the present time I am unable to deal exhaustively
with the matter, and, after a few remarks, propose merely tosupply some annotated
references supplementary to those furnished by Dr. Dalziel, and dealing almost entirely
with mosquitos known to be vectors of disease.

I hope that Mr. MacGregor, in charge of our Entomological Field Laboratory at
Wisley, in Surrey, may be able to carry out some research on the question in the
ensuing spring and summer, so far as it concerns indigenous species of mosquitos.

My attention was first specially directed to the subject when Mr. Harold King,
the Entomologist of the Wellcome Tropical Research Laboratories at Khartoum,
found Culex sitiens, Wiedemann (salus, Theobald), larvae in sea-water at Port Sudan, a
fact to which Dr. Dalziel refers in his paper, and when, at Khartoum, I found Anopheles
(Pyretophorus) costalis, Loew, breeding freely in brackish pools formed by seepage
through the “ weeping soil”’ of irrigation channels in Khartoum North. I had the
water of these pools analysed, but unfortunately cannot find the record. I think,
however, I am correct in saying that it contained at least 2 per cent. of common salt.

Hence this Anopheline can breed in waters similar to those in which, as Dr. Dalziel
mentions, Willcocks in Egypt found the larvae of Anopheles multicolor, Camboulin,
or, as it used to be called, Pyretophorus cleopatrae. This is of interest in view of
the fact that Graham recommended the salting of water containing the larvae of
A. costalis. He added common salt in the proportion of 3 per cent., and found that it
caused disintegration and precipitation of the motile algae upon which the larvae
feed. The latter, being thus deprived of their natural food, become cannibalistic.
Salt, he says, in lesser concentration appears to inhibit the growth of young larvae,
probably by diminishing their food supply, but seems to hasten the fully-grown larvae,
which become pupae more rapidly than usual.

As amatter of fact Dutton in 1902 had already shown that in the Gambia
A. costalis could breed in sait-water pools. Its larvae were found along with those of
Citlex thalassius, Theobald.

Apparently Grassi in Italy was the first to direct attention to Anophelines breeding
in sea-water. Nuttall, Cobbet and Strangeways-Pigg took the larvae of Anopheles
maculipennis, Meigen, in brackish water in England—twice in ditches, four times in
pools—and Christophers and Stephens discovered Anopheline larvae in brackish pools
at Accra containing 0-6 per cent. salt.

30 ANDREW BALFOUR.

Amongst the more important of the earlier investigations is that by de Vogel in
the Dutch East Indies. As Howard, Dyar and Knab point out, he disproved certain
conclusions of the Italian authors Perrone and Vivante, who stated that the maximum
proportion of sodium chloride in water which the larvae of Anophelines could resist
was under 2 per cent. Working chiefly with the larvae of A. vossi, Giles, de Vogel
found that a considerably higher percentage of sodium chloride, 7.e., as much as 2:88,
was not detrimental. His summary was as follows :—

“1. There are species of Anopheles which can live very well in sea-water.

“2. These mosquitos lay eggs which develop even in sea-water which has been
evaporated to half its original quantity.

‘3. These larvae in the gradually evaporating pools of sea-water can stand an
evaporation of the water to one-third of its bulk, but do not appear to
transform to adults if the concentration be greater than this.

“4. The larvae coming from eggs laid in sea-water of high concentration can
accomplish their entire metamorphoses in almost the normal time. This
is true even when the water has such concentration that the development
of larvae originally hatching in unconcentrated sea-water would be retarded
by this salt water.”

Later work by Carruthers and Christophers in the Andamans and also by Banks
in the Philippines and Swellengrebel in Java, suggests that de Vogel was probably
dealing with A. ludlowi, Theobald, rather than with A. rossi, the two mosquitos
being very similar.

A comparatively early record is that by Foley and Yvernault, who, in 1907, found
Pyretophorus chaudoyei, Theobald, breeding in Algerian waters of which the salinity
was greater than that of sea-water. P. chaudoyet is now called A. multicolor, an
Anopheline which comes very near A. /ispantola, Theobald, which, again, is probably
identical with the well-known Egyptian species A. turkhudi, Liston. As regards the
last-named, Gough has recorded the occurrence of its larvae in a brook of highly saline
water at Helouan, and succeeded in rearing them from water containing 2 per cent. of
salt. On the other hand Willcocks (loc. cit.) found that a 1 per cent. salt solution
proved fatal to the larvae of A. (Cellia) pharoensis, Theobald, a common Anopheline
of Egypt.

Banks in 1908 showed that in the Philippines 4. /udlowi, Theobald, a mosquito
mentioned by Dr. Dalziel in relation to the Andamans, breeds in both salt and fresh
water, and that in certain places they were present in water strongly impregnated
with lime and also containing much aluminium sulphate.

Clerc in France found the larvae of a Culicine, Ochlerotatus, (Culicada) cantans,
Meigen, in sea-water containing 44 grammes of chloride of sodium to the litre, and
recorded his observations in 1909. He noted that these larvae survive when trans-
ferred to fresh water, whereas, on the other hand, if the young larvae of A. maculi-
pennis are placed in salt water they die. The older larvae, however, survive and
become pupae and imagines.

In 1910 Gholap at Bombay discovered the larvae of A. stephensi, Rothwell, in
sea-water. The ponds in which they were found contained vegetable growths.

North American work on the subject, so far as Anophelines are concerned, includes
the observations of Chapin, who in Rhode Island found Anopheles larvae flourishing
in brackish coastal waters, and those of Smith on the larvae of A. quadrimaculatus,
Say, in the saline waters of New Jersey, and the records demonstrating the breeding
of A. crucians, Wiedemann, in similar localities both in New Jersey and Louisiana.

Stegomyia fasciata, Fabricius, figures in the list of mosquitos given by Dr. Dalziel
as breeding in boats and canoes, the water in which was for the most part brackish.

MOSQUITO BREEDING IN SALINE WATERS. oi

Hence it is of interest to note that in Somaliland Drake-Brockman recorded the
co-existence of Stegomyia larvae with those of Culex sitiens, and stated that brackish
well-water was acceptable to the former.

Macfie experimented with the larvae of Stegomyia fasciata at Lagos in the hope of
finding that common salt might be used as a larvicide in the case of this mosquito.
He showed that as regards domestic utensils the saline solution required to be of a
strength of 2 per cent. to ensure destruction of the larvae. Water containing 0-5 per
cent. of sodium chloride had no appreciable effect upon them.

Fielding, working at Townsville, Queensland, found that the female Stegomwia
fasciata oviposited in 70 per cent. sea-water.

Darling demonstrated the fact that in Panama Stegomyia fasciata could breed in
brackish water of varying chlorine content and that such water could also harbour
the aquatic stages of A. pseudopunctipennis, Theobald, A. malefactor, Dyar & Knab,
and Culex taentorhynchus, Wiedemann. In swamps containing 80 per cent. and more
of sea-water he found the larvae of A. albimanus, Wiedemann, and J. tarsimaculatus,
Goeldi, present in enormous numbers. Similar observations as regards 4. albimanus
were also made by Jennings of the Isthmian Canal Commission.

According to Peryassti the larvae of A. argyritarsis, Robineau-Desvoidy, in Brazil
are found in strongly brackish water as well as in fresh water.

Howard, Dyar and Knab mention that certain Brazilian observers, experimenting
with the larvae of A. albimanus and A. argyritarsis, “‘ found that in slightly brackish
water insagos were produced in a normal manner. In a mixture of 19 per cent. of
sea-water with fresh water only a very small proportion of larvae transformed to
imagos. Beyond this the larvae failed to pupate ; with 20 per cent. sea-water some
of the larvae survived three days ; with 30 per cent. all died after one day.”

Recent work by Taylor and Fielding in Queensland, Australia, shows that
A. annulipes, Walker, is occasionally found breeding in salt waters.

The latest research into the question of the occurrence of the larvae of Anophelines
in saline waters appears to be that by Sella, who had charge of the anti-malaria
campaign at Fiumicino, near Rome. His observations were carried out on the larvae
of Anopheles maculipennis or, as he calls it, A. claviger, and showed that a salinity up
to 6-7 per 1,000 of sodium chloride was without effect. Laboratory experiments
were also conducted, which indicated that the development of larvae is inhibited at
an early stage by concentrations of above 13-14 per 1,000, and that even those
exceeding 9 per 1,000 are no longer favourable to development. Pupae, however,
are able to tolerate a very strong concentration. From the point of view of the
employment of brackish waters in anti-malarial work it is interesting to note that the
investigations showed that solutions exceeding 20 per cent. must be employed in
order to ensure destruction within 10 hours, and exceeding 18 per cent. to obtain this
result within 24 hours. Further, it has been shown that larvae which have been
subjected to the action of the saline solution for several hours, if transported into
fresh water before they are dead, will survive. Apparently, to judge from cage

experiments, adult Anophelines are not influenced in laying by the mere salinity of
the water.

So far as Stegomyia fasciata is concerned, its behaviour as regards brackish waters
and sea-water has also formed the study of French and Brazilian observers, as recorded
by Howard, Dyar and Knab in their important and comprehensive work on the
mosquitos of North America.

Dr. Dalziel’s note on the various species of Culex found breeding in the coastal salt
marshes of the United States of America doubtless refers to the observations of
Chidester, who made an exhaustive study of the subject.

BP, ANDREW BALFOUR.

As regards the action of larvicides in saline waters the experiments of Jacob in the
Panama Canal Zone may be cited. Employing the well-known preparation devised
for use in that area and consisting of so-called “crude carbolic acid,” resin and
caustic soda, he proved by laboratory tests that the mineral salts in the salt water
do not lessen the toxicity of the larvicide. Moreover, if the larvicide be added as an
emulsion (and it is essential that this should be the case, for the sodium salts of the
cresols remain insoluble in sea-water), its entire quantity remains on the surface,
where it is found deadly to mosquito larvae. Jacob sums up to the effect that

“ When all other conditions are the same, the larvicide applied in an emulsified
state is more efficient in brackish water than it is in fresh water, and when
brackish water is to be treated, the area of the surface of the water is the
only factor necessary to be considered in determining the quantity of
larvicide to be used.”’

Perry, commenting on this work, stated that his experience in the field confirmed
the above results.

I have not been able to find records as regards the use of other larvicides in brackish
waters, but lack of time has prevented an exhaustive search of the literature, in which,
however, I believe there are but few references to the subject.

Mosquito larvae have been found in waters containing chemical constituents other
than the marine salts, for example, aluminium sulphate, as already mentioned. Some
of them, indeed, even survive in solutions which a priovt one would imagine should
speedily kill them. A consideration of these strange and sometimes sulphurous
nurseries would, however, lead us too far afield, but it might well form the subject of
another paper containing statements which, though perfectly true, might easily be
regarded as incredible.

References.

Baliour, A... 58 ss — Second Review. Supplement to
4th Report, Wellcome Tropical
Research Laboratories, Khar-
toumy 19Ut ips 20

Banks, C. S. sie .. A mosquito which breeds in salt Philippine Ji. Sci., Section B,
and fresh water. 1908, Vol. iu, No. 4, p. 335.

Carruthers .. 56 3 == Presidential Address delivered by
Surg.-Gen. C. P Lukis at the
second meeting of the General
Malaria Committee, Bombay,
loth Nov. 1911. ‘Abstract :
J, Drop: Med. ik srhys. WON
Vol. xv, p. 30.)

Chapin, C. V. ah .. The origin and progress of the Fisk Fund Prize Dissertation, No-
malarial fever now vrevaient 32. Providence, R.I. 1884.
in New England.

@hidester, HE. E: |... .. The influence of salinity.on the New Jersey Agric. Expt. Sta.
development of certain species New Brunswick. Bull. 299.
of mosquito larvae and its (Rev. in Rev. Appl. Entom.

bearing on the problem of the B, 1917, Vol. v., p» 103.)
distribution of species.

Chidester, F. E., & Patter- The influence of various concen- Entom. News, Philadelphia, vol.
son, R. trations of sea-water on the xxvii, No. 6, June 1916, p. 272.
viability of the salt-marsh (Rev. in Rev. Appl. Ent. B,
mosquitoes, Aédes sollicitans 1916, Vol. iv, p. 123.)
and Aédes cantalor (Dip.).

MOSQUITO BREEDING IN SALINE

Christophers, S. R.

Clerc, A.

Dalziei, J. M.

Darling, S. T.

De Vogel, W. T.

Drake-Brockman, R. E. ..

Dutton, J. E.

Fielding, J. W.

Foley, F. H., & Yvernault,

Gholap, R. D.

Gough, L. H.

Graham, W. M.
Grassi, B.

Howard, L. O., Dyar, H. G.
& Knab, F.

Jacob, J. E.

Kang, A. Ef.

Macfie, J. W. S.

(2416)

References—cont.

Malaria in the Andamans

Contribution a l’étude des mous-
tiques qui vivent dans !’eau
salée.

Crab-holes, trees, and other
mosquito sources in Lagos.

Studies in relation to malaria ..

Anophélines dans l’eau de mer.

Some notes on Stegomyia fas-
ciata in the coast towns of
British Somaliland.

Report of the malarial expedi-
tion to the Gambia, 1902.

Anophélines dans l’eau salée ..

Anopheles larvae breeding in

salt water.

Preliminary notes on Egyptian
mosquitos.

The study of mosquito larvae. .

The Mosquitoes of North and
Central America and the West
Indies.

The study of larvicides. I. The
action of larvicides in brackish
water.

Report on Economic Ento-

mology.

A note on the action of common
salt on the larvae of Stegomyia
fasciata.

WATERS.

Scientific Memoirs by Otficers of
the Medical and Sanitary De-
partments of the Government
of India, Calcutta, 1912.

C.R. Soc. Biol. 1909, Vol.
p. 120.

lxvi,

Bull. Entom. Res. 1920, Vol. xi,
Part 3, p. 247.

Isthmian Canal Commission
Laboratory Board of Health,
Department of Sanitation,
1910.

Atti della Soc. per gli Studi della
Malaria, 1907, Vol. viii, p. 1.

Ji. London School Trop. Med.
1913, Vol. ii, Part 3, p. 166.
(Rev. in Rev. Appl. Entom.
1914; Vol. ii, p. 72)

Liverpool School of Trop. Med.
Memoir No. 10.

Australian Inst. of Trop. Med.
Townsville, Queensland. Year-
ly. Report; jan. 1 toe Dec ot,
IOUS) ps de

Bull. Soc. Path. Exot.
Vol. 1, p. 172:

1908.

Ind. Med. Jl. June 1910, Vol. iv,
No. 6. (Abstract: Jl. Trop.
Med. & Hyg. 1910, Vol. xiii.,
p. 270.)

Bull. Ent. Res., September, 1914,
Vol. v, p.. 133.

Bull. Ent. Res., April 1910, Vol. i,
Pro:

Quoted by Howard, Dyar .and
Knab (q.v.), p. 228.

Volume i, 1912, pp. 224-229.

Proc. Med. Assoc. Isth. Canal
Zone, October 1912 — March
1913, Vol. v, Part)2, p86.

Third Report. Wellcome Re-
search Laboratories, Khartoum
1908, p. 206.

Bull. Ent. Res., February 1914,
Vol. iv, p. 339.

34 ANDREW BALFOUR.

References—cont.

Nuttall, G. H. F., Cobbet, Studies in relation to malaria. Jl. Hygiene, 1901, Vol. i, p. 4.
L. & Strangeways-Pigg, I. The geographical distri-
a bution of Anopheles in relation
to the former distribution of
ague in England.

Perrone, E. a6 .. Suicostumi delle larva delle zan- Atti Soc. p. g. Studi della Malaria,
zare del genere Anopheles in 1901) VolMiy pass:
relazione con le_ bonifiche

idrauliche.
Perry, we C- a .. (Discussion on Jacob’s paper, Proc. Med. Assoc. Isth. Canal
q.v.) Zone, 1912-1913, Vol. v, Bart
2, De 92s
Péeryassu, Aa G. 2s .. Os culicideos do Brazil. . .. Inst. de Manguinhos, Rio de
Janeiro, 1908.
Smith, J. B. He ab — Quoted by Howard, Dyar and
Kxnab (q.v.), p. 228.
sellay Vices. 36 .. The antimalarial campaign at Internat. J]. Pub. Health, 1920,

Fiumicino (Rome), with epide- Volsisp. oil:
miological and biological notes

Taylor, F. H., & Fielding, —— Australian Inst. of Trop. Medi-
VE we cine. Half-Yearly Report from
Ist July to 31st December1917,
p. 6.
Vivante .. ae 56 — Quoted by Howard, Dyar and
Knab (q.v.), p. 224.
WiillcockeH.(@o sa .. A preliminary note on the pres- Ann. frop. Med. & Parasit. 1910,
ence of mosquitoes in Cairo Vol. ili, p. 583.

and its environs.

ON SOME BORNEAN FIG-INSECTS (AGAONIDAE
—HYMENOPTERA CHALCIDOIDEA).

By JAMES WATERSTON, B.D., B.Sc.
(Published by permission of the Trustees of the British Museum.)

The following notes deal with the AGAONIDAE represented in a small collection
of fig insects formed in 1907-8, in Sarawak, by Mr. J. Hewitt, now Director of the
Albany Museum, Grahamstown, South Africa.

Genus Blastophaga, Grav.
Blastophaga jacobsoni, Grnd.

Blastophaga jacobsoni, Grandi, Boll. Lab. Zool. Portici, x, pp. 126 and 127, 26th
April 1916, and zbid. xii, pp. 21-32, figs. vii-x, 1917.

our 2°O “<Fromia- fig.”

BoRNEO : Siol, Sarawak, viii. 1908 (J. Hewitt).

In none of these specimens is the antenna complete beyond the sixth joint. Dr.
Grandi, who has at my request kindly compared this material with the type (which
was described from fruit of Ficus procera, Reinw., var. crassiramea, King, from plants
in the Botanical Garden, Buitenzorg, Java), is satisfied that it is referable to his
species, although the first mid-tarsal is not longer than the second, and the pilosity
of the thorax is not quite typical.

The range of Ficus procera and its var., so far as I can ascertain, is Java and
Sumatra. The species may, however; have been introduced into Borneo.

Genus Geratosolen, Mayr.
Ceratosolen hewitti, sp. nov.

°.—Head wider (10:9) than long (deep). Clypeal lobes large and very promi-
nent (fig. 1, a), central tooth short, keel distinct. Distance between the lower corner
of the eye and the base of the mandible slightly longer than the depth of the eye.
Antenna, length 0-9 mm. (fig. 1, 6) ; the scape viewed from beneath (fig. 1, c) and
outwardly shows a remarkable obliquely-set thickish edge or ridge ending abruptly
at both extremities ; process of the third joint (fig. 1, b, d) distinctly articulated,
fourth joint short ; sensoria of the funicle numerous, only shortly free distally, those
on the inner aspect of the fifth joint being slightly broader than any others ; spinose
bristles on the inner surface of the pedicel (fig. 1 d) numerous and stout. :

Trophi. Mandible with two small teeth and swollen along the anterior edge ; its
ventral surface with about five ridges. Appendage (2:1) short and broad, three-
fourths as long as the mandible itself and half as broad, with five ridges. Stipes
with one lateral bristle and no free splint.

Thorax. Protergum broadly concave posteriorly ; free striated margin broad,
narrower at the extremities ; whole sclerite bristly—40 bristles, more or less, on each
side of the mid line. Scutellum generally with three bristles along each furrow and
7-8 (minute) across the middle. Propodeon (fig. 1, e) with numerous bristles round
the spiracle. Prepectus showing a number (6-8) of fine striae, antero-posteriorly
directed, on the outer two-thirds. Mainly posteriorly, the mesosternum bears
numerous bristles.

Forewings twice as long as broad, length about 1-125 mm. Submarginal :
marginal: radius: postmarginal veins approximately in ratio 33:9:11:14. The
(2416) 2

36 JAMES WATERSTON.

submarginal bears towards the base a clear pustule (before which 1-2 bristles may
occur) without a bristle, but followed by five moderately long bristles, all on the
proximal two-thirds of the vein. At the uprise to the marginal is one somewhat long

Fig. 1. Ceratosolen hewitti, Waterst. sp. n., 9: a, edge of clypeus ; b, antenna; c, outline of
scape to show ventral lobe; d, armature of 2nd joint of antenna (pedicel), and details of
joints 3 and 4; e, right half of metanotum and propodeon ; f, abdominal spiracle.

C hewitti 3: g, head and thorax ; h, antenna.

bristle and 3-4 clear pustules triangularly arranged. Up to the radius the marginal
bears 7-8 bristles and the postmarginal 3-4; along the combined edges of marginal

BORNEAN FIG-INSECTS (AGAONIDAE—-HYMENOPTERA CHALCIDOIDEA). 37

and postmarginal stand about 20 bristles, some of which are double, 7.e., one rising
from above and the other from below the membrane. Radius with 4—5 bristles and
three clear pustules. Hindwing (7 : 2) 0-66 mm. long.

Legs. Fore coxa shorter (3: 4) than femur ; the latter (12 : 5) much longer (8 : 3)
than the tibia, which bears four teeth (alternately long and short) at the antero-
dorsal apical angle; tarsus approximately 4:2:2:2:5, the first joint with 2-3
bristles along the dorsal edge. Mid femur (4:1) longer (9:8) than coxa and tro-
chanter combined, with four longish bristles in a median row anteriorly and 1-2 more,
nearer the dorsal edge, apically ; posteriorly there are 8-9 bristles, mainly on the
apical two-thirds, 4-5 as on the anterior aspect being in a median row, with the others
above, towards the dorsal edge. Tibia distinctly longer than femur, narrow (10: 1)
on proximal two-thirds of its length, then rather abruptly expanding to the apex
(7: 1); along the dorsal edge are about 14 bristles (of which five or six on the apical
third are longer), and one or two fewer on the ventral edge ; additional bristles occur
distally on both faces, so that the apical third of the tibia is distinctly bristly. Tarsus
longer than the tibia, its first two joints in ratio 11: 7. Hind coxa externally naked, but
with about a dozen short bristles on inner surface—mainly on dorsal half ; femur (2 : 1)
not much longer (10: 9) than coxa ; tibia (12:5) rather broad towards apex. Tarsus
with first joint twice the second in length; chaetotaxy of first joint, dorsal 7-8,
anterior 12-13, ventral (plantar) 10—all beyond basal third—posterior 8-9 bristles ;
those on anterior aspect the thinnest ; second joint 3-4, 8-9, 5-6, 5; third joint
2, 6, 4, 3. The plantar spines are stout and rise from definite sockets, the plantar
edge not being frayed or fimbriated.

Abdomen. Tergites entire. The first (third) tergite bears over 30 minute bristles,
and the spiracle is enormous (fig. 1, f). Ovipositor barely projecting beyond the apex
of the abdomen. The length visible varies, according to pressure and relaxation of
the parts, from 0-04-07 mm. in a series of mounted examples. The last sternite
is truncated, except for a long, narrow median projection. Stylet (8 : 3).

Length, 1°5-1-9mm.; alar expanse, 2-8-3:4 mm.

g.-—Head length 0:45mm.; length of pronotum 0:35mm.; mesonotum
length 0-23 mm., breadth 0-4 mm; metanotum, length 0-175 mm., breadth
0:35mm.; propodeon, length 0-175 mm., breadth 0-35 mm.

Head (fig. 1, g) twice as long as broad anteriorly and about one-third longer than
its maximum breadth. The bristles (1, 1) set just above the median lobe of the
clypeus are short—half as long as their basal distance apart. Antenna (fig. 1, /)
with bulla about four-fifths the length of the scape (11:6); pedicel (4: 3) two-
thirds of the scape. Funicle (three joints) twice as long as the scape, its first joint
triangular in profile; fourth joint shorter (6:7) than fifth, but both greater than
the pedicel, which is only five-ninths of the fifth.

The propodeon is remarkably broad posteriorly (fig. 1, g).

Legs. Fore femur over twice as long as broad and more than two and one-third
times the tibia (excluding the apical tooth) in length ; tibia longer (10:7) than the
tarsus and not equal to the greatest breadth of the femur. Mid coxa, femur, tibia,
and tarsus subequal—the coxa very slightly the longest ; femur very convex dorsally
and subangulate at one-third from the apex ; apex of tibia with two teeth at upper
angle, one laterally and externally and two ventrally. Hind coxa (2:1), or with
membranous flange (10: 7), shorter (10: 11) than femur (4: 3) ; tibia (5: 2) not quite
three-quarters the length of the femur and as long as the tarsus, bearing 4-5 teeth
apically—counting both sides; tarsus sparsely set with bristles, one (lateral and
preapical) on joints 1-4 being longer.

Length, up to about 2mm.

38 JAMES WATERSTON.

Type 2 in the British Museum, one of a series of ¢ f and 9 9 from a fruiting
trunk of Ficus sp.

BoRNEO: Sarawak, vii. 1907 (J. Hewitt).

C. hewitti, sp. n., belongs to the group of which C. striatus, Mayr, and C. crassi-
tarsus, Mayr, may be taken as representatives. It is at once known in the 2 by the
short ovipositor and the terminal segments of the funicle, while the 3 is equally
characterised by the head and propodeon.

Genus Eupristina, Saunders.

Eupristina verticillata, sp. nov.

Q.—Head wider than deep (11:10); clypeus with a fine median ridge and
distinct but not prominent oral lobes, each with one bristle. The antennal grooves,
above the toruli, occupy about one-third of the width of the frons. Trophi: man-
dibles apically bidentate, with about five ventral ridges (fig. 3, a), anterior edge much
swollen ; appendage narrow, with 8-10 ridges (fig. 3, a, a1) of which 2-3 near the
articulation are strong, projecting tooth-like at the inner edge; maxillae (fig. 3, 5)
with no free splint. Antenna (fig. 2) just over 0-5 mm. long ; pedicel with a number

Fig. 2. Eupristina verticillata, Waterst., Span, e) : a, b, antenna; c, detail of joints 2-4;
d, apical sense-organs.

of stiff, rather fine, recurved bristles on inner dorsal aspect (fig. 2, c); process or
appendage of third joint long, completely articulated ; sense-organs on joints 5-11
long and tubular, subapical in position ; besides these on the outer apical edge of the
sixth joint a small cup-shaped sensorium with a short central process (fig. 2, b), and
another much larger one on the basal half of the eleventh joint (fig. 2 6, d); at the
extreme apex of the club a group of sensory spines—five in all (fig. 2,

BORNEAN FIG-INSECTS (AGAONIDAE—HYMENOPTERA CHALCIDOIDEA). 39

Thorax. Pronotum entire, but with a deep concave membranous area anteriorly ;
posterior edge gently and evenly concave, the free striate margin expanded abruptly
at the extreme sides into a subtriangular head ; spiracle not greatly salient. Medianly
and anteriorly the sclerite is bare, but at each side posteriorly towards the expansion
of the striate margin are 6-7 short bristles in two rows, and about a dozen more round
the spiracle. Scutum bare, except for one minute bristle on each side near the furrow
at one-third before the suture. Parapsides with six bristles. Scutellum and meso-

Fig. 3. Eupristina verticillata, Waterst., sp.n., 2 : a, mandible, ventral view ; a!, mandible
in profile; b, Ist and 2nd maxillae; c, wing; cl, pustules on submarginal vein; d, right half
metanotum and propodeon ; e, abdominal spiracle ; e1, the same, on same scale as d.

sternum each with bristles 3-4 : 3-4. Propodeon (fig. 3, d) with four moderate bristles
beyond the spiracle and one (occasionally two) minute near the posterior end of the
spiracle on the inner (admedian) aspect. Metasternum intumescent on each side of
the mid line, its surface raised in minute chitinous points.

Forewing (fig. 3, c, c!) just over 1 mm. in length, two and a quarter times as long
as broad, bare on about the basal fourth.

Legs. Fore coxa (2:1) with a longitudinal row of about 24 long bristles, which
form a fringe on inner aspect ; femur (17:8) about one-third longer than the coxa ;
tibia over twice as long as broad and equalling in length the breadth of the femur,
bearing at the apex antero-dorsally two teeth, with ventrally a short chitinous pro-
jection with one spinose bristle above; both anteriorly and posteriorly the tibia
bears a lateral row of about four short stiff bristles, and there are a few more along the

40 JAMES WATERSTON.

dorsal edge ; tarsus 2, 1, 1, 1, 3; the first joint bears one dorsal subapical bristle, one
anterior, one ventral, and three (stouter) posteriorly ; joints 2-5 have bristles 1, 1, 2, 1,
while five bears 1, 1, 0, 1. Mid coxa much widened posteriorly ; femur (10: 3)
shorter (6 : 7) than tibia (barely 6: 1), which has about eight short stiff bristles along
the dorsal edge and four ventrally on apical third. Tarsus as long as tibia, each
joint with one subapical bristle dorsally and another apically and ventrally ; joints
in ratio, 8, 7:7:5:9. Hind coxa (7:4) with 5-7 short heavy spinose bristles ;
femur (2: 1) one-seventh longer than the coxa; tibia (3: 1) three-fourths of the coxa
in length; tarsal joints in ratio 15:9:7:6: 11.

Abdomen. The tergites from the second (fourth) onwards are deeply incised
postero-medianly up to about one-half of their length. Between the spiracles is a
broadish membranous area, and this tergite is produced anteriorly into a broad,
angularly rounded median lobe. The stylet (8: 5) bears four bristles—three apically
and one at the side. The ovipositor sheath is slightly dilated apically, the ovipositor
extending beyond the apex of the abdomen a distance of about 0:6 mm. The spiracle
(fig. 3, e, et, same scale as d) is oval, flattened on one side.

Length (excluding ovipositor) 1-4 mm.; alar expanse, 2-4 mm.

Type @ in the British Museum, one of a small series of 9 9 “ From large tree

with small figs in middle of Museum grounds.”
BorNnEO: Kuching, Sarawak, x1.,1907 (J. Hewitt).

E. verticillata, sp. n., is an isolated form which I have placed in this genus with
some reluctance. It is easily recognised by its antennae, wings, protergum, etc. The
material available for description is in a very broken condition, there being only two
complete antennae and one wing in a score of specimens. Such dealation and loss

of appendages are commonly incurred when the female enters the fig in which the
eggs are to be laid.

The neuration of the single wing preserved is peculiar and perhaps abnormal.
There is a distinct marginal and postmarginal length, and in the membrane itself a
disconnected pellucid thickening, suggesting an obsolescent radius. Particular atten-
tion is for this reason directed to fig. 3, c, which illustrates the points referred to.

THE BIONOMIGS, OF TABANUS APREPES, AND OTHER
AUSTRALIAN TABANIDAE.

by Gi rite, FES,
Entomologist, Australian Institute of Tropical Medicine,
Townsville, Queensland.

(Plates I and IT.)

During recent years the number of described species of Australian TABANIDAE,
or March-flies, as they are almost universally called in this country, has been in-
creased very considerably, and with this increase there has come a wider knowledge
of the distribution of the various genera and species. While advancement has taken
place in this direction, little progress appears to have been made towards acquiring
an accurate knowledge of the life-history and habits of any of these flies, and, up to
the present time, no precise information has been published concerning oviposition,
larval development, feeding habits and pupation of any of our numerous species.

In this paper I propose to describe in some detail the life-history, habits and
developmental stages of Tabanus aprepes, Taylor, and T. rufinotatus, Bigot, which
have been reared from egg to adult, and to discuss other species so far as present
knowledge permits.

Tabanus aprepes, Tay].
I. aprepes, Taylor, Proc. Linn. Soc. N.S.W. xliv, p. 56.
IT. batchelort, Taylor, loc. cit. p. 58.

Distribution. This species is recorded from South Queensland (Eidsvold), North
Queensland (Townsville and Kuranda), and the Northern Territory (Darwin and
Batchelor).

Breeding-places. The following notes are based on observations made during the
period 18th October 1919 to September 1920, in three localities within the municipal
boundaries of Townsville, namely, (1) a small permanent rock-hole and stream arising
from it, situated within half a mile of the Institute building, (2) a small shallow
swamp about 250 yards distant from the rock-hole, and (3) a group of similar
swamps on the outskirts of the town.

During the wet season and for some months afterwards the rock-hole and
adjoining pools and riffles (Plates I and II, fig. 1.) are completely swept by a rapid
torrent of surface and soakage water from adjacent land; but later, when the flow
is reduced to a regular and steady trickle, there is present a plentiful supply of algal
growth and three or four feet of clear water in the rock-hole and rather less in the
pools. From June to December 1919 the moist banks were searched unsuccessfully
for larvae and pupae resulting from eggs deposited during the previous summer.
On 10th October 1919 and later, TABANIDAE were occasionally seen flying about,
or momentarily resting upon the heads of children playing on the sea beach distant
about 300 yards from the rock-hole—then the only locality within a mile or more
from which they could have emerged. The species could not be determined with
certainty, but was provisionally referred to T. aprepes.

More frequent visits were now paid to the locality in the hope of definitely associat-
ing with it the flies occasionally noticed in the vicinity. Digging operations were
undertaken wherever the soil was sufficiently moist or loose enough to permit of the
emergence of the flies from the ground ; then the harder and drier parts of the bank
and neighbouring soil were dug over and sifted, the rocks and herbage overhanging
the water were searched frequently for ovipositing females or their eggs, and a few

42 (Gees te SHOE IL

horses and cows which drank at the lower pools were watched, but no evidence of
TABANIDAE could be found until 6th January, or two days after the first heavy fall
of rain (106 points) since the preceding March.

On 6th January a female T. aprepes was observed on a twig about four and a
half feet over the water (Plate I), apparently about to oviposit, when she was
disturbed by a spider and flew off.

On 13th January three egg-masses were found on the same twig, all of which
appeared to have hatched, but on removal to the laboratory three living larvae were
rescued from a spider’s web which enveloped one end of one mass. Although there
appeared to be no unhatched eggs in the masses, the latter were placed over
water, and during the night 110 larvae were produced, some of which were subse-
quently reared to the fly stage and identified as T. aprepes.

From 16th January until 16th April numerous larvae of T. aprepes were found
in the algae floating on the surface of the water. Heavy floods swept the holes in
the locality during the month of April, after which larvae were not found.

On 2nd April a female T. aprepes was observed to alight on the twig from which
eggs were taken on the 13th January, and the process of oviposition was observed,
with the aid of a magnifying glass, from commencement to conclusion, when the fly
was captured and the eggs removed to the laboratory, where larvae were subsequently
reared.

Of the 39 egg-masses collected here (Locality 1) between 13th January and 9th
April, five were found on twigs and six on grass leaves or seed-heads overhanging
water in the rock-hole (Plate I), the majority being from 3 to 44 feet above the
surface ; three were found on grass leaves a few inches above the water trickling from
the rock-hole, and 25 on the terminal shoots of couch-grass overhanging a sloping
bank (Plate II), the surface of which was oozing with soakage water and algal growth.

The adjacent swamp (Locality 2) dried early in July 1919, and remained in this
condition until 4th January 1920. Towards the end of April 1920 the surface area
reached its maximum, there being then about 44 feet of water in the deepest parts.
Repeated searches were made for egg-masses on plants’ overhanging the margin and
on lily and other leaves floating on the surface, but none were found. During these
searches numerous half-grown to full-grown larvae (T. aprepes) were found clinging
to the lower surface of the lily leaves, or to the stems, or hidden in floating masses of
algae, from 20 to 30 yards from the margin and in from 3} to 43 feet of water. <A
few of these larvae were bred to maturity to confirm identifications.

The small lily-pond (Locality 3, Plate II, fig. 2) dried late in July 1919, and
remained so until 4th January 1920. In May of 1919, when the water was two or
three feet deep, a careful search was made for Tabanid larvae amongst the reeds
and water-lilies, but none were found.

From 10th June 1919 onwards, the banks of this and adjoining ponds were examined
from time to time as they dried, but in none of them were the larvae or pupae of
T. aprepes found, although other species were taken.

From January 9th to April 16th 1920, when the ponds and swamps contained
water, egg-masses and larvae were unsuccessfully sought for on many occasions.
On the latter date a careful search was made of the vegetation growing near the
banks and of the lily leaves in deeper water (4-5 feet) w ‘ith the result that many
larvae of T. aprepes, in all stages of development, were captured. On the same
date a fly of this species was observed ovipositing (1.45 p.m.) on the underside
of a seed capsule of a plant growing in eight inches of water and twelve inches from
the bank. When about 30 eggs had been laid, a lamp chimney, closed at one end with
mosquito netting, was slipped over fly and plant and tightly plugged with wadding.
The stem was then cut off about an inch below the wadding plug, leaving the

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 43

fly and eggs practically undisturbed. In this position the chimney and its contents
were returned to the water and overhanging plants. Several times during the next
hour the fly appeared to be on the point of continuing oviposition, but each time
returned to the netting without having extruded any eggs.

Between 21st and 25th April very heavy falls of rain and high winds caused an
accumulation of drift (grass, aquatic plants, twigs, cow and horse dung, etc.) to be
thrown up on the sloping banks of this and adjacent pools. When examined on 28th
April, numerous half-grown to full-grown larvae of T. aprepes, T. rufinotatus and
T. nigritarsis were found in the drift and on or under the soil beneath it ; while others
were found buried in the grass-covered soil between the outer fringe of the drift and
the foot of a stiff loamy bank three or four feet from it (Plate IT, fig. 2). The latter
were full-grown and were in some cases obviously at rest in the positions in which they
intended to pass through the long dry period to follow; in others they were still
burrowing downward. Quite a number of those found in the drift, and especially in
that part near the water’s edge, were evidently feeding, as shown by the contents
of the alimentary tract. Many of these larvae (TI. aprepes) were captured and bred
out in the laboratory in July and August.

While turning over the soil and drift on this date several recently discarded pupal
cases were found and subsequently identified as those of T. vifinotatus and T. nigritarsis.
These pupae were almost certainly the product of eggs laid more than a year earlier,
since the unusual conditions of the 1919-20 season and the shortness of the possible
breeding period (103 days) almost preclude the possibility of their being derived from
eggs laid during the current year. No definite evidence has been obtained to determine
the maximum period during which the larvae of these insects may remain in a dormant
condition, but certain facts suggest that full-grown larvae may, after the usual long
resting period and in the face of a drought, postpone their transformation into pupae
for six months or more.

On the same date (28.iv.1920) and a few yards distant, full-grown larvae of T.
aprepes were found making their way up a slope from the water’s edge, through
short wet grass and litter towards a bank (similar to that shown in Plate II, fig. 2)
and about eight feet from the water’s edge. Some were actually travelling when
observed, others were sheltering in the grass or under debris.

These banks were examined four months later (30th August), when several T.
aprepes (three pupae and five larvae) were found, and on the same afternoon an adult
female was captured in the vicinity. The period (winter) intervening between these
dates, 28th April-30th August, had been unusually mild and moist, doubtless favouring
early development and rendering the emergence of the adults possible after even
a light shower of rain. During the preceding year, owing to drought conditions,
emergence of adult flies from this soil would have been a physical impossibility
throughout the whole period April 1919 to January 1920.

Oviposition. In discussing breeding-places brief reference has been made to two
flies which were observed in the act of ovipositing on 2nd and 16th April. As the
process differed in several respects from that recorded for Indian and American
species, it may be of interest to refer to it more fully here. At about 4.30 p.m. on
2nd April, whilst sitting by the water’s edge (Locality 1), a fly was seen to alight
on the underside of a twig about four feet above the surface of the pool (Plate I)
and about three feet from my head. The twig lay at an angle of about 45° to the
water, and the fly settled on it head uppermost. After lightly touching the surface
of the twig many times with the tip of the abdomen, the first egg was extruded whilst
only the extreme apex of the abdomen was flexed. Immediately the apex of the egg
touched the twig it appeared to stick, and as it left the body it was forced out of the
vertical position to an angle of about 20°; the apex of the abdomen was again applied
to the twig at the point of attachment and to one side of the egg just laid and a second

44 (515, 1s0NbIl,

extruded in the same manner. Three eggs were then laid close to and in front of the
first two, each one as it left the ovipositor being pressed firmly against those behind,
until the latter lay almost parallel to the twig. In this way three or four gradually
widening rows were laid, each slightly nearer vertical than the preceding one. Then
a second and similar tier was commenced on top of those already laid, the first row
of eggs of the second tier resting on the second or third row of the lower tier. Moving
her body slightly forward, the fly extended the lower tier two or three rows before
continuing the upper tier. In this way the operation was continued until the
mass had covered the lower side of the twig for a length of 30 mm., when the fly
moved forward a few millimetres and remained stationary until captured. The rate
of oviposition was about three eggs a minute, and at no time was the apex of the
abdomen brought under the thorax, as stated by writers dealing with extra-Australian
species.

The Egg-mass. The size and shape of the egg-masses are very variable. As a
rule they are about 20-33 mm. long by about 2mm. wide when deposited on slender
twigs or narrow blades of grass, or they may be more compact when laid on seed-heads
or other objects offering a wider base for the mass. The mass is invariably as wide
as the object upon which it rests and generally contains two layers of eggs. Some
of the masses, however, contain three layers of eggs and are proportionately shorter
and higher. Sometimes a second and even third mass is laid very near to or over-
lapping part of an earlier one, and small masses containing only a few score of eggs
are found near masses of average size. The number of eggs per mass probably
averages about 500, but much smaller and much larger masses, 7.e., 250-700, are
found. At first the mass is creamy, but in the course of about 24 hours it changes
to light slate-purple, and gradually darkens with the development of the larvae.
There is a complete absence of the white chalky substance used by some species as
an outer coating of the mass, and more often than not its general appearance is
distinctly rough and lacking finish.

Fig. 1. Tabanus aprepes, Tayl., eggs. Fig. 2. 7. aprepes, embryo 78 hours old.

The Egg. The eggs measure 1-60 mm. to 1°65 mm. in length, by 0-3 mm. in
width, and are variable in shape according to their position in the mass, but always
bluntly rounded at either end (fig. 1). The surface is smooth and glossy and, in
recently laid eggs, pearly white. The colour changes rapidly ; eggs that were laid at
1.50 p.m. on 16th April changed to grey by 5.15 p.m., to dark slate-purple by
9.30 a.m. on 18th, and hatched during the night of 22nd or early morning of 23rd.
Another batch of eggs which was laid at 4.30 p.m. on 2nd April changed to
dark slate-purple by 9.30 a.m. on 4th and hatched between 2 and 4 p.m. on 8th.
The egg period therefore was from six and a quarter days to seven days in the one
case and about six days in the other.

For at least 24 hours before the young larvae hatch they are distinctly visible
through the clear, thin shell (fig. 2). During this period they are active, dark-banded
little creatures, with the head always directed outwards from the point of attachment

THE BIONOMICS OF AUSTRALIAN TABANIDAF 4

of the egg-mass. When about to liberate themselves the blade-like process (egg-
cutter) at the anterior end is pressed against the apex and drawn downwards along
the side in two or three deliberate strokes, which cause a rent in the shell sufficiently
large to permit the young larva to escape rapidly.

Larval Development. Generally speaking, there is only an interval of a few minutes
between the appearance of the first and last larvae from a given batch of eggs. Some-
times, however, the interval is much longer, and this is especially the case with
large masses, in which, owing to their form, many of the eggs are concealed beneath
tiers of other eggs. One large mass from which all the larvae had apparently emerged
before noon (13th January) produced 110 additional larvae before 9 a.m. on the
following morning; another produced about 600 larvae during the morning and 30
more late in the afternoon.

In only one instance (5th April) was the dispersal of larvae observed under natural
conditions. On this occasion several very small larvae were gathered on the surface
of the pool (Plate I) in one dip of the scoop while searching for Anopheles, and upon
examining some twigs overhead an egg-mass was found from which larvae were then
dropping. The mass contained approximately 300 eggs, arranged in a single layer
eight or ten abreast, of which number about one-third had already hatched and
dropped into the water ; the balance emerged during the succeeding five or six minutes.
Upon reaching the water the young larvae were rather sluggish, but sufficiently active
to disperse by slow lashing movements, some remaining on the surface film, others
seeking the shelter of floating masses of algae. Some of these larvae were collected
and removed to the laboratory for examination and observation.

In the laboratory the egg-masses were usually suspended over a shallow dish of
clean water, into which the young larvae dropped as they freed themselves from the
mass. For some hours they remained more or less quiescent on the surface, during
which period the first moult was accomplished. This process commences before, or
immediately after, the young larvae leave the egg, and is sufficiently advanced to be
seen under a low power five minutes later. Apparently the first moult is always
completed during the first six hours of larval life. As a number of larvae from one
egg-mass have been reared through all their stages to the perfect fly, the development
of these may be now recorded. While this batch was under observation a number of
other batches were available for study, and they supplied much information and
material for examination which could not have been obtained from the constantly
decreasing ranks of the original batch.

The egg-mass from which the larvae were obtained was found partly evacuated
on 13th January (Locality 1); 110 young larvae emerged from it on the afternoon
and night of 13th-14th January and 105 of these comprised the original batch.

First Instar. An accurate description of the young larva soon after it emerges
from the egg is somewhat difficult, owing to the fact that ecdysis has already com-
menced. When about five minutes old and after fixation in the usual way the young
larva measures about 1-40 mm. long by 0-28 mm. wide at the sixth segment ; the
segments appear to be withdrawn into each other; the cuticle about to be cast off
envelopes the body loosely, excepting at the head, where it is firmly attached. The
surface is pale in colour and distinctly marked with longitudinal striae; several
moderately long slender hairs are present about the middle of each segment ; there
are no short spine-like hairs fringing the anterior margin of the first three segments
(thoracic), such as occur on the fourth segment and in increasing numbers on the
fifth to the tenth segments. The anterior margin of the first two segments and the
anterior and posterior margins of the following seven segments appear to be banded,
but this appearance is due to characters on the cuticle beneath. The mandibles are
withdrawn into the head, but in cleared specimens they are seen to be short, curved
rods arising apparently in the anterior third of the first segment. The egg-breaker

46 Gb. Hike.

is a black, chitinous, angular projection of the upper anterior margin of the labrum
and is the most conspicuous feature of the first instar. Graber’s organ is visible on
the posterior portion of the tenth segment as a pair of dark, pyriform, closely
approximated bodies.

Second Instar. Since most of the characters found in the larvae of the second
instar are visible through the cuticle of, and appear to belong to, the first instar, the
changes which follow the first ecdysis do not appear to be so great as they are in reality.
Fixation in hot alcohol after the first ecdysis is completed and in all subsequent stages
of larval development has the usual effect of extending the body to its fullest length ;
thus, whereas the length of a larva of the first instar is about 1-40 mm., the length
just after moulting is about 1-00 mm. greater. The measurements, descriptions and
figures which follow are all from specimens fixed in hot 70 per cent. alcohol, and
therefore fully extended. The characters of the larvae of the second instar are shown
in fig. 3, which represents a larva between one and six hours old. Such larvae range

Fig. 3. T.aprepes, anterior end of larva 1-6 hours
old (above) ; posterior end of same (below).

from 2-35 mm. to 2-63 mm. in length, by 0-375 mm. to 0-425 mm. in width at the
widest part ; the segments bear distinct longitudinal striae ; the anterior border of
each segment, excepting the first three (thoracic), is fringed with numerous small
bristle-like hairs and dense short brown pile, which produce the banded effect (seg-
ments four to nine inclusive resemble the tenth segment shown in fig. 3). The first
three (thoracic) segments are shown in fig. 3. The mouth-parts are prominent ; the
first maxillae, palpi, and antennae are easily distinguished ; the anterior projection of
the labrum is rounded and no longer black and chitinous; four malpighian tubes are
distinguishable ; Graber’s organ still contains one pair of bodies, which are now seen
to be enclosed in a pyriform sac, the rounded end of which is foremost ; the apical
third of the syphon tube bears two groups of three long slender hairs, which usually
appear to arise at the extremity owing to invagination of the apex and withdrawal
of the stigmal plate.

During the 16 or 17 hours following the first ecdysis there is little apparent change
in the young larva other than a slight increase in size and, generally, the addition of
another pair of bodies in Graber’s organ. A long series of larvae of this age measured
from 2-35 mm. to 2-82 mm. in length, by 0:35 mm. to 0-425 mm. at the widest part.

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 47

For about three days following their emergence the young larvae do not feed, but
remain on the surface film. After this period, however, they commence to attack
each other, even in the presence of an abundant supply of small shells and other
animal life introduced upon aquatic plants.

When twelve to fourteen days old the majority of the larvae measured from
6 mm. to 7 mm. in length, by 0-7 mm. to 0-85 mm. in width ; the groups of hairs on
the syphon tube increased to four or five, and Graber’s organ contained either two or
three pairs of bodies, each pair diminishing in size from the anterior end. A few of
the larvae had developed very slowly during these twelve days and now measured
only 4-5 mm. in length. In two larvae (7 mm. and 8-5 mm. in length respectively)
a second pair of mandibles could be distinguished in the anterior third of the first
segment, and faint traces of the dark bands characteristic of older larvae of this
species could be detected near the junction of the segments and near the anal
protuberance. The second ecdysis apparently takes place when the larva is between
7mm. and 9mm. in length. Unfortunately the number of larvae in this batch
became so reduced, largely owing to cannibal practices, that specimens could not be
secured as frequently as desired to determine this point, but a “ wild” larva captured
on 2nd March appeared to represent an early stage of the third instar and to connect
the last-described individuals with older and more advanced ones of the same batch.
This larva measured 9mm. in length by 1-5 mm. at the widest part ; the junctions
of the segments were banded and blotched with dark brown, and the dorsal tubercles
and the pseudopods bore short bristle-like hairs, as in older larvae.

On 2nd March, or when 49 days old, two larvae of the original batch measured
15mm. and 16 mm. in length respectively. Whether the subsequent changes in the
larvae follow ecdyses, or whether they are developed gradually during the third instar,
has not yet been satisfactorily determined. On 16th April, or when 94 days old,
two other larvae of this batch were destroyed by their fellows. The former now
measured 21-5 and 24 mm. long by 3 and 4 mm. wide respectively. The brown bands
and blotches are now very distinct and of the same.pattern as in adult larvae: the
striae are well marked on all segments, but are absent on the brown areas; the man-
dibles are black ; the bunches of stout curved spines above the insertion of the antennae
are pale ferruginous, very prominent, and overhang what appear to be moderately
large facetted eyes situated behind them; in the smaller of the two larvae there are
five pair of bodies in Graber’s organ, in the larger six pairs ; the stigmal plate is now
visible at the apex of the syphon tube. About the anterior third of the first segment
(prothoracic) there are several long branched hairs, on other segments they are simple
or absent ; the pseudopods are prominent ; there are no hairs on the anterior margin
of the thoracic segments, these being confined to the dorsal ridges and pseudopods
of the abdominal segments, where they are inconspicuous. The dark bands and
blotches are formed entirely of very short and dense pile, as in the younger larvae ;
the pseudopods were very prominent in the larger of the two larvae.

On the same date (16th April) the contents of the breeding-tray were collected,
with the object of separating the remaining larvae of T. aprepes and several larvae
of TL. mgritarsis which had been placed in the tray recently. The former now
numbered only five, two of which measured about 28 mm. long and three about
33 mm. long. One of the latter was retained for examination and is described below, the
others were put separately into five-inch flower-pots half full of moist, clean granite
sand, upon which lily leaves and shells were placed every three or four days. The
larva referred to above was 94 days old at the time of its death, and measured 33 mm.
in length and 5mm. in width. In size and coloration, as well as in external form, it so
closely resembles the mature larva which has undergone a long resting period that a
description of it will suffice for both. There are six prominent pseudopods to each
segment from the fourth to the tenth inclusive, arranged on the anterior margin, three
on each side of the median line; each pseudopod is armed with a number of short
bristle-like hairs ; on the dorsal surface of the same segments there are two transverse

48 Ga ny Weel.

tubercles situated on the anterior margin on either side of the median line, which
appear to be of the same nature as the pseudopods and similarly armed. The dorsal
surface is blotched and banded boldly with dark brown (fig. 5), the pattern of which
is very characteristic and constant. The ventral surface of segments four to nine |
inclusive are creamy white; from the dark anterior margin of the first segment

Fig. 6. T. aprepes, adult larva, anterior end.

Fig. 7. 7. aprepes, adult larva, posterior end.

(prothoracic) there extend posteriorly five dark, lanceolate marks, one on either side
of the dorsal and ventral surfaces, and one in the median line of the ventral surface.

The second and third segments are banded anteriorly (fig. 6) and the tenth posteriorly
(fig. 7) with brown.

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 49

As the larvae reached maturity and ceased to feed (about the third week in April)
they were placed together again in a large receptacle containing moist sand, in which
they yemained for about two months. At first they were very active, coming
to the surface at night and returning to the sand during the day, but never attacking
each other. On 3rd June they were all in a torpid state and contracted to about
18 mm. in length, but when placed in water they regained their normal appearance
and activity.

On 25th June one of the larvae, now 164 days old, pupated in a vertical position
four inches below the surface and after a pupal period of 11 days emerged as a perfect
fly (female). Another larva pupated about a week later and was preserved as a
specimen ; the third pupated on the 7th July and produced a male fly on 21st July,
or 190 days after hatching; the fourth remained in the larval stage until 8th
September (239 days), when it was destroyed for examination.

Larval Habits under Natural Conditions. Under natural conditions the larvae are
to be found commonly on or near the surface of clear and moderately deep pools,
amongst submerged herbage near the banks, resting upon the lower surface of lily
leaves, upon the submerged stems and leaves of all kinds of aquatic plants and in
floating masses of algae. Clear and moderately deep water appears to be essential.
The food of the young larvae is not known, but it is believed to be the small molluscs,
which were fed successfully to larvae reared in captivity. Several kinds of molluscs
are very plentiful in most of the breeding-places, and these certainly form an important
part of the diet of older larvae, which have frequently been found feeding on them.

In the rock-hole (Localitv 1) this food is absent, but there is a plentiful supply of
other kinds. Cannibalism is common amongst ‘‘ wild”’ larvae, and is practised by
individuals in all stages of development. To give some idea of the voracy of these
insects, it may be mentioned that on one occasion 12 large larvae were placed in a
pickle-jar of water and algae for transportation to the laboratory—a journey of
three miles—and upon arrival there only two remained alive, one of which destroyed
the other before arrangements could be made for their separate accommodation.
On another occasion over 50 nearly full-grown larvae were placed in a large
porcelain dish with sand, water and fresh molluscs. Within four days many of
the latter and 40 of the larvae were destroyed.

With the aid of a small stout wire net attached to the end of a long bamboo rod
the plant growth can be disturbed sufficiently to dislodge the larvae, which, even if
carried down by the currents thus created, soon appear near the surface and are easily
captured in the net. In some cases a stout wire hook at the other end of the rod was
found useful for dragging masses of vegetation towards the bank for closer inspection.
Wading was resorted to in many cases, but the results were usually unsatisfactory on
account of the restricted range of vision.

The larvae appear to live entirely in water until they have reached maturity,
when they migrate from the water to high ground close by, as described elsewhere in
this paper, and penetrate into the soil or clay to a depth of from 7-15 cm., where they
remain in the larval stage, generally with head uppermost, for several months.

The Pupa. During the first few hours following metamorphosis the entire pupa
is buckthorn-brown,* but the eyes soon deepen to mummy-brown, and then to blackish
brown, while the thorax becomes argus-brown. The average size is about 21 mm. in
length by 4 mm. in width at the thorax, the Ist and 6th segments slightly narrower
than the 2nd to 5th inclusive.

On the dorsal surface of the first abdominal segment there are two stout hairs on
either side of the median line and three on each pleura, two of which arise close
together near the wing-sheaths. Segments 2 to 7, inclusive of both surfaces, bear an

* Ridgway’s colour nomenclature.
(2416)

50 Gob. BILE:

uninterrupted fringe of long and short bristles arranged roughly in two rows, the
shorter bristles nearer the base ; on each segment from the 2nd to 5th these bristles
increase in length ; on the 5th, 6th and 7th they are equally long or, rarely, the long
bristles may be absent from the middle of the seventh tergite and sternite and the
shorter ones may be branched.

The six projecting spurs at the apex of the abdomen are arranged three on each lobe
of a prominent bilobed tubercle divided vertically by a deep cleft, which is wider in the
males ; the upper and middle pair are equally long and stout, the lower are smaller ;
on either side of the dorsal surface midway between the base of the tubercle and the
apex of the 7th segment there is a group of four or five stout spines of unequal length.
The length and stoutness of these bristles vary greatly in individuals of either sex.

The anal tubercle is very large and deeply furrowed in the male and is bordered
anteriorly by an unbroken fringe of about 18 to 22 stout bristles of variable lengths.
In the female the fringe is broadly interrupted in the middle, and consists of from 6-9
bristles on either side (fig. 8).

Pig. 8. 7. aprepes, ventral surface of last segment of female pupa.

Duration of the Pupal Stage. The duration of the pupal period during the months
June-September varied from 8 to 20 days, the average being 12 days in the case of
10 individuals whose periods were accurately observed.

The factors which determine early or late emergence of laboratory specimens are
not known, and no explanation can be offered at present of the fact that whilst certain
pupae of a batch pass through a very short pupal stage—e.g., eight days—-others of
the same batch and subjected to the same treatment may remain in the stage for
periods up to 20 days.

Emergence of the Fly. The emergence of laboratory-bred flies has been observed
on several occasions—always between 10 a.m. and 4.30 p.m.—and in each the pro-
cedure was similar. The pupa, with its body vertical, works its way to the surface,
from which it protrudes the thorax and first two or three abdominal segments. In
this position it remains motionless for from one to two hours; then, with very little
apparent effort, the thorax is burst on the dorsal surface to the posterior margin of
the mesothorax, and also along the lower margin of the eyes, leaving a flap-like piece
bearing the anterior group of tubercles more or less loosely attached ventrally. The
fly frees itself in a minute or two, leaving the greater part of the pupal case as before.
At emergence the wings are uniformly opaque, but in the course of an hour or two
they become hyaline or suffused with brown, as the case may be.

Ihe Adult. The adults of this species show considerable variation in regard to
size and coloration and, as might be expected, in laboratory-bred specimens this is more

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 51

apparent than in those bred under natural conditions. Inarecent paper* it has been
stated that the wings show various gradations in colour from a total absence of
infuscation to a very pronounced shading, especially at the veins. A long series of
males and females bred out in this laboratory show these gradations to a marked
degree, not only in flies reared from full-grown larvae captured in natural surroundings,
but also in flies reared from the same batch of eggs. Variations in abdominal colora-
tion of both sexes, and especially in the females, is even more pronounced. In some
specimens there are evident spots at the apex of certain segments, in others the
abdomen is of uniform dark brown or uniform ochraceous tawny. In three wild
females captured in June in this district the colour of the abdomen is so light as to
lead the writers of the above-mentioned paper to refer them with some doubt to this
species. Recently, however, a similar form has been bred from a batch of eggs which
also produced individuals showing all the variations of wing and abdominal coloration.
It would appear that in this and other species too much attention has been paid in the
past to slight or even marked differences of this kind. When these variations in
colour are associated with abrasions of the abdomen and thorax, and especially of the
front, the possibilities of error in diagnosis are greatly increased unless a fairly long
series 1s available for comparison.

In life the lower third and the lateral and posterior margin of the eyes of the male
are claret-brown with emerald-green iridescence ; the upper two-thirds are drab grey
and composed of coarse facets. In the female the facets are small throughout and
the colour uniform drab grey with brown iridescence.

Seasonal Occurrence. Throughout Australia, perhaps excepting the wet belts of the
North Queensland coast, of which I have little knowledge, there is a marked seasonal
occurrence of flies of this family, namely from about October to April. In the southern
regions, where there are regular winter rains and low winter temperatures, the
emergence of flies is probably regulated by temperature and not by rainfall; but in
the north, where the temperature is more or less equable and the rainfall is, excepting
for moderate falls, confined to the summer months above mentioned, the condition of
the soul, whether dry or wet, is most certainly the regulating factor. Thus almost
invariably the march-fly season is either early or late, good or bad (from the collector's
point of view) according to whether the summer rain is early or late and normally
heavy, light or absent.

The chief natural breeding-places in this district, and in most others with which
I am familiar, are in temporary accumulations of water, such as shallow swamps,
which disappear completely in the winter or dry season and after the full-grown larvae
have entered the soil in the vicinity of the water’s edge, wherever that may be
at the time of their reaching maturity. As the soil dries these larvae are practically
immobilised, and remain so until it again becomes thoroughly moistened. It
follows, then, that there can be no emergence of flies until after heavy rain has fallen.

The truth of these statements has been demonstrated on many occasions, and
notably during the summer of 1919-20, when until 4th January the country was in
the grip of a severe drought, and there was an almost complete absence of Tabanids
in this district. Heavy rains for a short period in this month were followed by the
appearance of a moderate infestation of flies. (It may be stated here that the previous
season also was @ne of unusually low rainfall, and therefore an unfavourable one for
fly breeding.) The January rainfall was not sufficiently heavy to raise the level of
the swamps to normal, and another long spell of dry weather intervened before the
banks in the vicinity of the normal high-water marks were thoroughly saturated.
About the middle of April abundant rains filled the swamps, and there followed a
marked increase in the number of flies in the district. Whether the flies were derived
from eggs laid in January 1920 or during the preceding wet season could not be

* Ferguson & Hill, in the press (Proc. Linn. Soc. N.S. Wales).
416) DZ

52 Gib, HIE

determined, but the period required for the life-cycle, as ascertained later, suggests
strongly that the latter was more probable. Further, there is no evidence to suggest
that the Tabanids in this or other districts known to me produce two generations in
one season. Obviously these remarks do not apply to localities in which there is
permanent surface water, such as Locality No. 1.

In some localities a few individuals of certain species, particularly T. rufinotatus,
are to be found throughout the dry season (June to August), but their occurrence can
be accounted for by the presence of permanent water (such as Locality No. 1), or at
any rate marshland, in the vicinity.

During June of this yearnumerous young larvae were found in pools which dried
a few weeks later. The question naturally arises whether these larvae perish or
whether they are able to burrow into the mud to lie dormant through the remainder
of the dry season and until conditions again become favourable for their development.
Attempts to determine these questions have been unsuccessful, but it may be mentioned
that one larva from a late batch of eggs was, when about 9 mm. long, accidentally
isolated in the laboratory in a pot of dry sand and without food of any kind from about
Sth June to 28th August. On the latter date it had the general appearance of adult
larvae in the resting stage preceding pupation, and became active immediately it
was placed in water containing suitable food and cover. Unfortunately neglect
caused its death before any development could be detected. Hine, working in
America, found that certain Tabanid larvae after a long resting period again fed
before pupating, but nothing of the kind has been observed to take place in the case
of any of the species studied here.

Rearing Larvae in Captivity. Various methods of rearing the young larvae were
tried, but as none of them gave satisfactory results it is not intended to describe them
in detail. In order to prevent cannibalism young larvae were isolated in small earthen-
ware pots with a capacity of about 120 c.c., prepared in various ways, and supplied
with various kinds of food. Apart from the difficulty of keeping the water in these
vessels fresh, this method was found to be too cumbersome, and the larvae rarely
survived for more than a week or so. Larger vessels, 7.e., five-inch flower-pots and
small museum jars, were equally unsatisfactory. Kerosene tins cut lengthwise into
two equal parts gave better results, and in them a few larvae were reared from
the egg to maturity. These tins were prepared by placing a quantity of clean sand
at one end and two or three inches of water at the other. Pieces of water-lily
leaves, algae and swamp plants carrying small molluscs were placed in the water
to afford shelter and food, and these were renewed as often as possible. The water
was changed every four or five days by lifting one end of the tin and allowing it
to filter through the sand. The tin was then partly filled and again emptied in
this way, before being finally replenished with water and food-bearing vegetation.
Each tin contained the progeny from one batch of eggs, 7.e., 300-600 larvae, of
which never more than 1-2 per cent. reached the imago stage. When molluscs
were not obtainable mosquito larvae and small earthworms were offered as
substitutes, but worms were invariably refused by larvae in all stages of their
development. Mosquito larvae were destroyed by the Tabanids when the former
were stranded in algae or sand, but otherwise they appeared to have been able to
avoid capture. At all times Tabanids of this species appear to feed upon their fellows
in preference to molluscs or any other animals, and for this reason it was found best
to transfer the survivors, when of nearly full size, to separate pots or dishes, where
they completed the development in moist sand, upon which molluscs and portions of
lily leaves were placed. The best results were obtained from a batch of 105 larvae
which were reared in a concrete trough, measuring 16 inches wide by 20 inches long
by 6 inches deep, prepared similarly to the tins just referred to. As the larvae
approached maturity the water was gradually reduced until it was confined to a small
area at one end of the trough. In this molluscs were placed every week or so, until

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 53

it was found that none were being devoured, when the water was drawn off and the
trough half filled with clean sand, which was kept moist. From time to time the sand
was turned over to watch the progress of development, but beyond this the larva
received no attention for intervals of several weeks. Some of the larvae pupated and
_ produced flies in this trough, but most of them were transferred to small pots of clean

moist sand, where they completed their development. In future it is intended to use
these large troughs or galvanized iron trays instead of the smaller vessels.

Tabanus rufinotatus, Big.

T. vufinotatus, Bigot, Mem. Soc. Zool. France, v, 1892, p. 673.
T. lineatus, Taylor, Rept. Aust. Instit. Trop. Med. 1911, p. 65.
T. elestéem, Summers, Ann. Mag. Nat. Hist. (8) x, 1892, p. 224.

T. designatus, Ricardo, Res. Expéd. Sci. Néerlandaise Nouv. Guinée, ix, pt
1913, p. 390.

Distribution. This is a widely distributed species, having been recorded from
South Australia, New South Wales, Queensland and the Northern Territory, and also
from Dutch New Guinea. The South Australian specimens in the British Museum
collection are most probably from Port Darwin (Northern Territory) or its vicinity,
which prior to 1911 formed part of the state of South Australia.

Breeding-places. On 25th March 1919, an egeg-mass was taken from the lower
surface of a Juncea leaf growing three feet from the bank of a shallow water-hole
(near Locality 3) in twelve inches of water. At the time the pool contained a
maximum depth of three feet of water and was much frequented by cattle and
horses which grazed in the district. As the season advanced the water dried back
rapidly and finally disappeared before the end of May. During the period interven-
ing between 25th March and 28th May 1919 the vegetation near the water and the
muddy banks and bottom were searched for egg-masses and larvae respectively,
but without success. During the same period and up to the present date (September
1920) many other possible breeding-places have been repeatedly examined, but so
far only a few adult larvae have been secured.

The Egg-mass. The egg-mass referred to above, from which the larvae described
in the following notes were derived, measured 5-5 mm. in length by 4:25 mm. in width
at the base and 3-5 mm. in height. The eggs, which numbered about 500, were
arranged very compactly in three tiers and were coated, either separately or collec-
tively, with a white secretion. After the larvae hatched, the mass retained its form
so perfectly that, viewed from any direction but from above, it appeared to be still
composed of viable eggs.

Larval Development. The larvae hatched almost simultaneously at noon on 30th
March and moulted between that hour and 9 a.m. on the following day. Specimens
were not secured prior to ecdysis, so that the first instar cannot be described here.

The Second Instar (figs. 9, 10). When 24 hours old the larvae of the second instar
measured from 2:5 mm. to 2-6 mm. in length by about 0-25 mm. in width at the
widest part. Up to this age they were of uniform size, but at the end of the second
day variations became apparent, some measuring 3 mm., while others had increased
to3-S5mm. The cuticle is creamy white and bears minute longitudinal striae; the
tufts at the base of the antennae are short and many of the component hairs are forked ;
the prothoracic segment is short, and the hairs which fringe its anterior margin and
the anterior margin of the next segment are difficult to discern. The dorsal tubercles
and pseudopods are armed with moderately large hairs, and each segment bears
several isolated long, slender, pale hairs.

The young larvae remained in a small dish of water for three days without food,
when they were placed in a large concrete trough containing sand at one end and

54 G, Fo Air

water at the other. In this they were offered small molluscs, young mosquito larvae
and small aquatic animals that adhered to the water-lily leaves used to provide
cover. From time to time dead larvae were found which appeared to have been
destroyed by their fellows, but the remainder seemed to thrive. On 10th June, or
when 77 days old, one of the larvae measured 15 mm. in length. Unfortunately this
specimen was lost before a detailed description was obtained. On 3rd July the sand
was allowed to dry off gradually, and from this date onwards small earthworms only
were offered as food. -On 8th October the sand was washed over, but no larvae were
found. The vessel and its contents were left undisturbed until 27th October, when
the now dry sand was sifted, yielding three larvae measuring from 22 mm. to 23 mm.
in length by about 2°5 mm. in width. In such larvae the cuticle is glossy and devoid
of the striae observed in larvae of the second instar, there is no trace of banding, and
the hairs on the dorsal tubercles and pseudopods are colourless.

Me
~

WA atc

re
Hise

|
|

Sedan?

hr
if

a x

Fig. 10. T. vufinotatus, posterior end of larva 24-48 hours old.

The two remaining larvae were now placed in a large shallow dish, containing sand
at one end and water at the other, in which they were fed on earthworms and mosquito
larvae until 8th November, when they were placed separately in five-inch flower-pots
standing in a dish of water and containing three inches of clean sand. On 8th December
both larvae were evidently fully grown. One of these was secured for examination
and is described below, the other was returned to its pot, to which had been added a
few small earthworms and a little earth. A week later it was found dead and damaged
beyond recognition by scores of nematode worms. The loss of the sole surviving
larva rendered the identification of the species with which I had been dealing for over
nine months impossible for the time being, but the larvae secured on 27th October

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 55

and 8th December provided the material from which a determination was

subsequently made in comparison with a series of “ wild ” larvae, some of which were
subsequently bred out.

The Adult Larva (figs. 11, 12). The full-grown larvae measure from 26 to 29 mm.
in length by about 4 mm. in width, and are creamy white in colour, with the faintest
indication of darker bands; the cuticle i is glabrous and without striae; the tufts at
the base of the antennae are composed of short stout hairs ; there are no hairs on the
anterior margin of the thoracic segments, and those of the dorsal tubercles and
pseudopods are very short, slender and dark in colour.

Fig. 11. 7. vufinotatus, anterior end of adult larva,

Tig. 12. TV. vufinotatus, posterior end of adult larva.

From this record it will be seen that under laboratory conditions the larval stage
was not less than eight and a half months.

Development of Larvae under Natural Conditions. Larvae were taken on 28th
April 1920 in association with the larvae of T. aprepes and T. nigritarsis (see notes on
T. aprepes), and again in a similar bank on 15th June in association w ith the larvae
of Silvzus notatus. After securing specimens for examination, the remainder were
placed in small earthenware vessels containing sand, which were thereafter kept moist.
The flies emerged as follows :—one female on 93rd May (after a pupal period of nine
days), one male on 2nd June, and one female on 21st July.

The Pupa. The average length of the pupa is about 15 mm. and the width at the
thorax about 4mm. The eyes are blackish ; the vertex (in the female) and thorax
Dresden brown, the latter with faint traces of stripes; abdomen ochraceous tawny,
with Dresden brown stripes corresponding to the black stripes on the abdomen of the

56 G. By BibT,

imago. The first abdominal segment bears two isolated hairs on either side of the
median line and one on each pleura behind the spiracle; the second to seventh
tergites and pleurites each have a double row of bristles, those of the anterior row
being much shorter than those of the posterior. On the ventral surface, segments 2,
3 and 4 each bear an irregular single row of bristles of varying length; on
the Sth to 7th segments the bristles are arranged in two rows; the terminal
segment bears the usual number of large spurs (six) ; anterior to the anal tubercle
there is, in the male, a fringe of about twenty long straight or curved bristles,
which in the female is interrupted in the middle and is composed of five to eight stout
and generally straight bristles (fig. 13). In the male the anal tubercle is large and
deeply ribbed, in the female it is small but prominent. On either side of the dorsal
surface, midway between the base of the upper pair of spurs and the posterior margin
of the 7th segment, there is a group of six or eight stout bristles of irregular size and
shape, which in the female is reduced to two or three much smaller bristles; in
addition, in the male, there is on either side a group of two or three very short bristles
midway between the above and the posterior extremity of the anterior fringe.

Fig. 13. ZT. vufinotatus, ventral surface of last segment of female pupa.

Lhe Adult. In life the eyes of the male have the upper part grey, with deep brown
iridescence, and the lower part, lateral and hind margins maroon, with an emerald-
green band in line with the insertion of the antennae. The facets of the lower part
are larger than those of the upper. In the female the eyes are maroon to dark maroon-
purple, with two broad green bands in line with the callus and subcallus respectively.

Seasonal Occurrence. These flies were very scarce indeed during the period
February 1919 to September 1920. Females were seen or captured in the field in the
months of January, February, April, June, September and December, generally
while buzzing about one’s head or resting upon one’s hat. Horses (locally and in the
Northern Territory) appear to be more troubled than cattle, the former usually being
bitten about the nose, ears, rump and coronet.

Tabanus nigritarsis, Tay].

T. nigritarsis, Taylor, Rept. Aust. Instit. Trop. Med. 1911, p. 67.

Distribution. This species has been recorded previously from North Queensland
(Houghton River) and the Northern Territory (Darwin, Stapleton, etc.).

Breeding-places and Habits. Between the 19th May and 10th June 1919 five
apparently full-grown larvae were gathered from the submerged stems or leaves of

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 57

various plants growing in a few feet of water at the margin of a small swamp (Locality
No. 3, Plate II) and removed to a concrete trough containing a pile of sand at one end
and water at the other. In this vessel they were fed upon molluscs until 3rd July,
after which date food was refused by them. The water was now drained off and the
sand allowed to dry gradually until only the bottom two inches remained moist, in
which condition it was kept for about four months. On 19th August one of the
larvae was found in a vertical position about three inches below the surface of the
sand. The body was then much contracted, but when placed on the surface
the insect became more or less active and extended its length to about 25 mm.
This specimen was allowed to bury itself again, and remained undisturbed in the
sand until 8th October, when it and two other larvae were found in the above
condition. One of these was preserved in the usual way and is described in these
notes ; the others were placed separately in five-inch flower-pots full of sand and
imbedded in the sand contained in the larger vessel, where the two remaining larvae
of the original batch of five were presumed to be resting. The two larvae in pots
remained very active, but refused to eat either molluscs or earthworms, although they
came to the surface almost nightly until 18th November. On 26th November one of
these larvae was found undergoing metamorphosis, the other following on the morning
of Ist December (11 a.m.), the former producing a female fly on 12th December and
the latter a male on 17th December (between 9 and 10 a.m.).

On 27th November the sand in the large trough was sifted and another larva
secured, which pupated on 12th December (5 p.m.) and produced a female fly on 28th
December. The fifth larva was not recovered and was, therefore, presumed to have
been destroyed by its fellows during the early days of their captivity. From the fore-
going it will be seen that the pupal period in the laboratory was 16-17 days. Through-
out the greater part of their lives in captivity these larvae and pupae showed much
restlessness, and in the latter days of their pupal existence frequently wriggled to the
surface or projected the posterior end above it, proceedings which would have been
impossible under natural conditions owing to the hardness of the soil in which they
would have been embedded.

As previously noted (under 7. aprepes) a few larvae of this species were found in
the resting stage in a clayey bank in this locality on 25th April 1920, associated with
T. aprepes and T. rujfinotatus. On 15th June following five additional specimens of
T. nigritarsis were taken under similar conditions in the same locality. At the time
of writing (30th September 1920) the majority of these are still in the larval stage in
moist sand, others which at the time of capture were embedded in balls or tubes of
plastic clay and placed on the laboratory shelves to dry were equally healthy and
active when released on 30th September.

The Larva. The adult larva measures about 35 mm. in length by 5-5 mm. in
width and is cream-coloured, faintly blotched and banded with yellow ochre (figs.
14-17). The pseudopods are arranged two on either side of the median line on the
ventral surface and one on each pleura, these and the dorsal tubercles being only
slightly elevated. The surface of the cuticle is dull and distinctly marked with
longitudinal striae, except where banded or blotched.

The Pupa. The pupa measures about 23 mm. in length by 4 mm. across the thorax
and 5 mm. across the abdomen at the widest part. The eyes are blackish brown, the
thorax slightly lighter and the abdomen argus-brown. The first abdominal segment
bears two slender hairs on either side of the median line, one on each pleuron behind
the spiracle, and two closely approximated hairs on the ventral surface near the margin
of the wing-sheaths. On the dorsal surface of the 2nd segment there is a double
row of bristles, the anterior row short, stout and of irregular size, the posterior much
longer; the 3rd segment is similar to the 2nd; on the 4th, 5th and 6th the bristles
increase in size gradually from the 4th posteriorly ; on the 7th there are fewer long
bristles in the posterior row, their place being occupied by others of intermediate size.

58 (in, 1000.

On the pleurae the armature is similar to that of the dorsum, except on the 7th segment,
which resembles the ventral surface. On the ventral surface the Ist, 2nd and 3rd
segments bear a single row of mixed short and long bristles increasing in length from
the 2nd; the 5th and 6th segments resemble the corresponding segments of the

Wig. 17. 7. nigritarsis, posterior end of adult larva.

dorsum, the 7th is similar, but has some short bristles in the posterior row. The
integument of both surfaces bears transverse striae on the anterior two-thirds of each
segment, which are generally absent or replaced by punctures on the posterior
third. The armature of the terminal segment is variable. In most specimens there

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 59

is on the dorso-lateral margins, and midway between the base of the second pair of
projecting spurs and the posterior margin of the 7th abdominal segment, a well-
developed group of from five to seven very irregular bristles arising from a raised base.
In some specimens these bristles are reduced to three or four in number, and in one
they are absent on one side and on the other represented by two rudimentary bristles.
In the male the anal opening is bordered anteriorly by a semicircle of stout and very
irregular bristles, some of which are expanded and branched at the base, others short
and almost rudimentary, or all may be long and moderately slender. Posteriorly the
opening is bordered by a raised and deeply furrowed prominence. In the female
(fig. 18) the fringe of bristles in front of the anal opening is broadly interrupted in the
middle, each group being composed of from five to eight bristles of variable size.

The Adult. In lite the eyes of the male are dull yellow-green and coarsely facetted
above the point of junction of the eyes on the front ; below this and on the sides and
posterior margin the facets are smaller and bronze-coloured, with gold and green
iridescence. In the females the eyes are uniformly dull yellow-green.

Fig. 18. T. nigvitarsis, ventral surface of last segment of female pupa.

Seasonal Occurrence. Apart from the three flies reared in the laboratory only two
individuals (females) have come under my notice during the period 25th February
1919 to 30th September 1920. Both were captured whilst attacking horses during the
last week in February of this year. In the Northern Territory (1912-1917) this
species was regarded as the most numerous and most troublesome to stock during the
months of December-February. The egg-masses found there were small and
compact and were generally placed on the underside of pandanus leaves overhanging
water.

Silvius notatus, Ric.
S. notatus, Ricardo, Ann, Mag. Nat. Hist. (8) xvi, 1915, p. 264.
S. psarophanes, Taylor, Proc. Linn. Soc. N.S.W. xlii, 1917, p. 520.
S. fuliginosus, Taylor, op. cit. xl, 1915, p. 810.

60 GSEs HILT.

Distribution. This Tabanid is a widely distributed species, having been recorded
from S.W. Australia, Victoria, New South Wales, South Queensland, North Queensland
and the Northern Territory (S. fuliginosus, Taylor).

Although so widely distributed, it would appear to be a rare species in this district,
where, apart from those referred to above, only one specimen (the type of S. psaro-
phanes) has been recorded hitherto. Nothing is known of its early stages and the
feeding habits of the larvae and adults.

Breeding-places and Habits. On 22nd August 1919, while breaking down and
sifting the low banks of a small lily-covered pool (Locality No. 3) about 24 Tabanid
larvae were found in nearly dry, stiff, loamy soil at depths below the surface varying
from 10 to 30 cm. In most cases the larvae lay in a vertical position with the head
uppermost, others lay horizontally, and a few vertically with the head downward. At
the time the bank was distant 12 feet from the water’s edge and six feet from the mud
which intervened between it and more or less dry soil. All the larvae were in the lower
part of the bank in soil which had been above water-level since about 12th May, and
although a search was made in the mud and adjacent dry soil no larvae were found
there. The undamaged larvae, 16 in number, were removed to the laboratory and
placed in two five-inch flower-pots containing moist soil, in which they remained undis-
turbed until 8th October. From the 22nd August until 22nd September they were
very restless and wandered over the surface of the soil at night. These movements
ceased on the latter date, and on the 8th October about one inch of the surface soil
in one pot was temporarily removed, exposing the head and thorax of three pupae.
The colour of the eyes and wing-sheaths indicated that they were then from about
three to five days old. The oldest was preserved as a specimen, and the others
were placed separately in pots of earth, from which they emerged as flies (3 and 9)
on 13th October after a pupal period of about eight days. On the same date two
males emerged from the second pot. Other flies emerged as follows:—1 3 on 25th
October, 1 g on 3rd November, 1 2 on 8th November, 1 3 on 24th November,
1 2 on 26th November, 1 Q 30th November, 1 3 2nd December (pupal period 14
days), 1 9 12th December.

On 15th June 1920 two resting larvae were taken from the bank shown in Plate II,
fig. 2, and were transferred to small pots of sand, where they still remain in the larval
stage (30th September).

Lhe Larva, The adult larva measures about 33mm. long by 4mm. wide and is
of a creamy white colour with narrow orange citrine bands at the anterior end of
each of the first ten segments. On the first three segments the banding is obscure and
on the 11th it is confined to a collar-like expansion of the posterior margin. Each
abdominal segment, excepting the last, bears a slightly elevated transverse ridge or
tubercle about half as wide as the segment, as well as a pseudopod on either lateral
margin and a pair of pseudopods on the ventral surface. Macroscopically the dorsal
and ventral surfaces are similar in appearance. The integument is glabrous and
bears very distinct longitudinal striae. The first and last segments, and especially
the spiracle (figs. 19, 20), differ greatly from those of any species of the genus Tabanus
known to me. Grabner’s organ has not been made out.

The Pupa. The pupa measures from 18-20 mm. in length, 3-3-5 mm. in width
across thethorax, and 3-5-4-0 mm. across the widest part of the abdomen. The colour
varies, according to age, from chestnut-brown to mars brown, head and apices of
wing-sheaths blackish brown, lower surface and sides of first abdominal segment
ochraceous tawny. In the male the head is as wide as the thorax, in the female
slightly less. The abdomen is nearly cylindrical in both sexes. The thoracic spiracles
are large and overlap the posterior margin of the head. The first abdominal segment
bears two slender hairs on the tergite and one on each pleurite near the wing-sheath.
The second tergite bears an interrupted single row of very short stout and irregular
bristles and three or four long bristles, generally towards the sides. The third, fourth

THE BIONOMICS OF AUSTRALIAN TABANIDAE. 61

and fifth tergites are similar, except that the bristles are stouter and there are about
eight long bristles on each. On the sixth and seventh tergites the bristles are fewer
and stouter than on the preceding ones. On the pleura they are arranged roughly
in two rows, those in front being much shorter and fewer than those behind. Sternites
2-7 are armed similarly to their corresponding pleurites. On the anterior half of each
tergite there are five or six small dark-coloured depressions, the foremost being
nearest the pleura and the hindmost nearest the median line. Behind the anterior
margin of each segment and parallel with it there is another row of three or four
similar depressions on either side of the median line. The anterior two-thirds of each
tergite, pleurite and sternite are distinctly marked with transverse striae ; posterior
to the bristles these striae are less distinct or absent and the whole surface is punctate.

Fig. 19. Silvius notatus, Ric., anterior end of adult larva.

Fig. 20. Szlvius notatus, posterior end of adult larva.

The terminal segment is deeply rugose, the two upper projecting spurs or bristles
are parallel along their inner margins in the female and divergent in the male ; the
_middle pair is much larger than the upper and lower and projects laterally. The
anal tubercle of the male is very large and deeply furrowed ; in front it is bounded
by a continuous fringe of bristles, four or five comparatively small ones in the middle
and a group of four or five very stout ones on each side, similar to but larger than

62 G. F. HILL.
those of the female (fig. 21) ; the latter group of bristles is continued, with a slight
interruption, around the sides of the segment to a point in line with the base of the
lower pair of spurs. The size and number of these bristles are variable in both sexes,
but are generally largest and fewest in the males.

Fig. 21. Szlvius notatus, ventral surface of last segment of female pupa.

The Adult. In life the eyes of the female are of uniform light seal-brown and the
facets of equal size throughout. In the male the greater part of the surface of the
eye is of the same colour, but at the lower third it is crossed by a sinuous, iridescent
blue-green and copper band, which extends to near the lateral margins and is con-
tinued (in light seal-brown) around the posterior margin to the vertex. The facets
forming this band are very small, while the remainder are large.

Butt. ENT. RESEARCH. VoL. XII. Part 1. Prane: Ik

Rock-hole (Locality No. 1), showing spot upon which eggs of Tabanus aprepes,
Tayl., were laid (marked: with a cross).

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BuLi. ENT. RESEARCH. VoL. XII. Part 1. PLATE II.

Fig. 1. Pool in stream (Locality No. 1); twenty-five egg-masses of Tabanus
aprepes were found on grass overhanging the bank.

Fig. 2. Clay bank (Locality No. 3) from which larvee of Tabanus aprepes,

T. rufinotatus, T. nigritarsis and Silvius notatus were obtained.

THE LIFE-HISTORY OF EUTHYRRHINUS MEDITABUNDUS, FABR., AN
IMPORTANT WEEVIL PEST OF MANGO TREES IN AUSTRALIA.

By G. F. Hit, FES.
(Plate III.)

The adults of the weevil, Euthyrrhinus meditabundus, Fabr., were frequently
_ taken in the forest country to the south of Darwin, Northern Territory, and on several
occasions they were bred from the branches of two species of indigenous trees and from
custard apples, Poincianas and two unidentified species of introduced ornamental
trees growing in the Darwin Botanic Gardens. As a rule the infestation of these
trees was not heavy, although Poincianas occasionally showed considerable injury.
Mango trees, which were frequently badly damaged and sometimes destroyed by
termites (Mastotermes darwinensis), appeared to be immune from attack by
Euthyrrhinus, even when growing in close proximity to infested trees of other kinds.

Later, while a resident of Townsville, North Queensland, I had opportunities of
studying the habits of this insect in the réle of a mango pest of the first importance.
Although these trees are not grown extensively for commercial purposes, they are
to be found in many North Queensland gardens, where they are prized not only for
their shade and beauty, but also for their prolific crops of fruit.

The following notes are the result of observations made on a number of large
mango trees growing in the vicinity of the Australian Institute of Tropical Medicine,
Townsville. In March 1919, two trees in particular attracted attention owing to the
number of dead twigs appearing through the dense foliage at the summit, and although
a close examination was not made then, the condition was attributed to the effects
of drought during the preceding dry season. During the dry season of 1919 (April to
the end of December) the condition of these trees gradually became worse, and others
in the vicinity showed symptoms of disease. Early in this year (1920) one tree
threw out bunches of leafy shoots along the main and upper branches, but the older
foliage continued to fall. At the end of April this tree died, and of the other there
remained alive only one secondary branch and a strip of bark extending from it to the
ground. From that month (April) to the end of August the beetles emerged in great
numbers from the trunks and main branches. Those beetles which emerged from the
now dead tree migrated at once, presumably to living trees in the vicinity, while those
from the other gathered on, and oviposited in, the remaining living portion.

Upon emerging from their holes the beetles are very active and begin at once to
seek mates. The act of copulation occupies about ten minutes, after which the male
remains attached to the female for some considerable time—two or three hours in
some cases. During copulation the females, and afterwards both sexes, gnaw away
the outer weathered surface of the bark and feed upon the living tissues or sap beneath,
leaving small areas of exposed tissue to mark the feeding places. This injury is
negligible, even when many beetles are confined in close captivity to a small
area of bark.

Oviposition.

The average period of development of the embryo within the ovary has not yet
been satisfactorily determined. On 6th August a pair were observed copulating at
noon, and an hour later they were removed toa healthy mango tree, on which they were
enclosed in a small wire-gauze cage tacked to the bark. During the following three

64 GE: HILL:

days the male was generally found clinging to the female, but copulation was not
again observed. The male died on the sixth day, and on the eighth day the female
was seen to have the proboscis deeply buried in the tissues of the bark, apparently
engaged in preparing a hole in which to insert an egg. Later in the day she was
attacked and crippled by small ants (Phetdole megacephala). On the following day
she was dissected, and an egg was found in the oviduct. Later two batches of six
couples each were confined on the bark of the same tree inasimilarmanner. Most
of the males died within ten days, and on 29th September there remained alive only
one female of one batch and three of the other. On the latter date the cages and their
contents were removed to fresh sites for further observation, and the bark previously
covered by the cages was carefully examined for eggs or larvae. Although the surface
showed numerous punctures, some of which extended to a depth of 3 mm., no trace
of eggs or larvae could be found. The reason for this apparent delayed oviposition
has not yet been determined, but it may be found to be due to refusal to oviposit in
captivity even under conditions so closely simulating nature.

At noon on 8th September a wild female was observed with the proboscis inserted
in the bark to a depth of about 3mm. At 5.30 p.m. the hole was found to be plugged
with comminuted bark, under which an egg lay 2 mm. from the surface. This egg
was removed in situ to the laboratory, where it hatched 7 days later. About this
time several other eggs were found in similar circumstances.

The egg is a rounded oval measuring 0-528 mm. in width by 0-688 mm. in length,
pearly white in colour, with a very finely granulated surface.

Larval Development.

Upon hatching the young larvae tunnel into the bark, subsequently, and while
yet very small, boring into the sapwood, obliquely or horizontally at first, but always
horizontally sooner or later. The duration of the larval and pupal stages has not yet
been ascertained, but it is believed to total 12 months at least. Experiments designed
to determine these points, andalso the length of life of the perfect insects, are in progress.

Appearance of Infested Trees.

The earliest observed symptoms of infestation are pronounced withering of the
terminal twigs, followed by partial defoliation, the development of numerous adven-
titious tufts of foliage (Plate III), and the death of the secondary and, later, the main
branches. The cause of these conditions is not manifest until the perfect insects begin
to emerge from the dead or dying branches or, very rarely indeed, from branches in
which there is still a flow of sap. As a rule the injury caused by the larvae is such
that the branch is destroyed before the first beetles emerge.

The bark of a heavily infested branch will be seen to be pierced by numerous
clean-cut holes (fig. 1) of various sizes, ranging from 1:5 mm. to 7 mm. in diameter,
the smallest of which, 7.e., those up to about 2 mm. in diameter, are made by the adults
of Hymenopterous parasites which have in the larval stage destroyed the young
' Euthyrrhinus larvae, the larger by adult weevils. The latter holes vary a good deal
in size in accordance with variations in the size of the adults, 7.e., from 2 mm. to 6:5
mm. in width by 4-5 mm. to 12-5 mm. in length. The males are generally, if not
always, very much smaller than the females ; but this statement appears to be often
contradicted by the finding of pairs of approximately equal size in a copulatory
attitude. If, however, these pairs are dissected they will be found to be females.

On the removal of the bark the sapwood shows a few shallow surface grooves made
by the young larvae before commencing their drive towards the heartwood, but more
often they bore straight in towards the centre without preliminary tunnelling. A
large proportion of the young larvae fail to penetrate deeper than 5 or 6 mm. before

THE LIFE-HISTORY OF EUTHYRRHINUS MEDITABUNDUS, FABR. 65

they are overcome by parasitic Chalcids. The survivors penetrate much deeper—
2.é., from 30-35 mm.—and when nearly full-grown turn about so as to direct the head
outward. In this position they enlarge the burrow to one uniform diameter from its
base to near the cambium, the frass accumulated in the process being very tightly
packed behind so as to fill from 10-22 mm. of the burrow, in the remaining portion
of which pupation takes place.

Fig. 1.—Main branch of a mango tree after the emergence of the weevils.

On emerging from the pupal stage the imago appears to undergo a rather long
developmental stage within the pupal chamber, during which the body hardens and
the insect becomes sexually mature.

The operation of boring its way out through the thin layer of sapwood and the
Lark which lies between the chamber and freedom is evidently a long and tedious
process, judging by the notches worn on the sides of the hole by the leg joints of the
toiling insect.

Parasites.

Early in August, when the beetles were first noticed to be plentiful, a species of
Chalcid wasp was frequently found in close association with them, or on adjacent
foliage. Selecting a branch from which Euthyrrhinus were then emerging and which
showed, in addition to their exit holes, a few smaller holes in the bark, two pieces, each
about 2 feet in length, were sawn off and removed toa cage, where during the next few
weeks about a dozen of the Chalcids emerged. On examination it was found that these
insects had developed in holes about half an inch deep, which had been made by the
young weevil larvae prior to being parasitised. Mr. A. P. Dodd, to whom specimens
were submitted, informed me that, although known to him from the Cairns district,
they belong to a new species, a description of which,under the name Chalcis euthyrrhint,
appears below. A few weeks later several specimens of Thaumasura curculionis,
Girault, were reared from the same branch and from the feeding-holes of young
larvae of the same host. About the same time both parasites were also captured on

(2416) E

66 GES ae

mango trees infested with Euthyrrhinus. It may be remarked here that the type
and other specimens of Thaumasura curculionis were reared by me in Darwin
(Northern Territory) from the branches of custard-apple trees and an undetermined
introduced tree infested with Euthyrrhinus meditabundus. A third and somewhat
similar Chalcid, of which a single specimen only has been secured, is also a parasite of
this weevil. Mr. Dodd has identified this species as Thaumasura pavo, Girault.
In addition to the above, two specimens of a much smaller parasitic wasp were bred
from the eggs of this host, while a rather brightly hued Braconid has been taken fre-
quently on infested branches. Although the latter has not been definitely associated
with E. meditabundus as a parasite, there is little doubt as to its relationship with the
latter.

It is only too evident that these parasites have not proved themselves to be of
much practical value in controlling this pest, and that artificial means must be resorted
to if efforts are to be made to prevent the destruction of these valuable shade and
fruit-producing trees.

Artificial Methods of Control.

No experimental work has been attempted with the object of devising means for
combating the ravages of this insect, a pest which will probably prove difficult to
control, and for several reasons. The adults are capable of flight as soon as they
emerge from their burrows in the wood; and certainly disperse soon after emergence
from trees which do not provide suitable conditions for rearing future broods. For
this reason badly infested trees should be cut down and burnt before the adults
leave them, for it is evident they cannot be saved once the main branches and trunks
show evidence of heavy infestation. Whether the spread of the beetles from infested
to healthy trees could be effectively checked by this means remains to be proved, but
there is some evidence to suggest that such a measure would have the effect of localising
the area affected.

Peane lil

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PART

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Bult. ENT. RESEARCH,

Mango tree, showing effects of attack by a weevil,

Euthyrrhinus meditabundus, Fabr.

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A NEW CHALCID PARASITE OF EUTHYRRHINUS MEDITABUNDUS.

By ALAN P. Dopp.

Chalcis euthyrrhini, sp. nov.
9.—Length, 5-7 mm.

Head and thorax black ; abdomen bright orange-red ; tegulae red; antennae
clear orange-red, the two apical club joints darker, the scape darker and varying to
black ; posterior legs, including the coxae, clear orange-red ; anterior and intermediate
coxae black, their femora and tibiae more or less brown or fuscous, their tarsi red.

Head, viewed from above, transverse, the occiput concave and (from lateral aspect)
declivous, immargined ; frons gently yet distinctly convex from eye to eye, viewed
from in front distinctly wider than deep ; antennal scrobes long, narrow, reaching to
and hardly containing the median ocellus at base, divided by a wedge-shaped elevation
that tapers to a point at half their length ; eyes large, bare ; ocelli large, in a slightly
curved line, the lateral pair separated from the median ocellus by less than their own
diameter ; cheeks broad ; surface with dense uniform umbilicate punctures. Antennae
separated at base, inserted slightly above a line drawn across ventral end of the eyes,
13-jointed, counting the club as 3-jointed ; scape slender, as long as the next four joints
combined ; pedicel not niuch longer than its greatest width ; ring-joint large, nar-
towed at base ; funicle 1 twice as long as its greatest width, 2 slightly longer than
wide, 7 a little wider than long ; club conical, fully twice as long as its greatest width,
the first suture distinct and slightly before the middle, the second suture faint.
Thorax normal, sculptured like the head, the sculpture very coarse, without con-
spicuous pubescence ; pronotum large ; its posterior margin deeply concave, the scutum
thus much produced at meson anteriorly ; anterior margin of scutum with dense
scaly sculpture, rising sharply from the pronotum ; scutellum hardly longer than its
greatest width, its apical plate rounded ; axillae rather widely separated ; propodeon
almost perpendicular in relation to the scutellum, moderately long, coarsely rugose
and also finely shagreened, with a shallow median channel, laterally and near anterior
margin with a distinct subacute tooth or projection on either side ; pleurae sculptured
like the rest of the thorax, the mesopleural depression longitudinally striate. Fore-
wings ample, normal, stained yellowish, venation black ; marginal vein three times
as long as the stigmal, which is as long as the post-marginal. Abdomen, viewed from
above, pointed ovate, fully twice as long as its greatest width, the valves of the
ovipositor slightly exserted ; gently convex above and beneath, no longer than the
thorax ; segment 2 (first body segment) almost as long as the others united, 3 longer
than 4, 5 and 6 subequal and shortest, 7 fully as long as 4-6 combined, 8 rather short
and with a median carina ; 2 smooth and polished ; 3-6 smooth, except for yellowish
pubescence against their anterior margins, no setae at meson, the free area broader
on 3, narrower on 6 ; 7 with rather dense shallow punctures and yellow pubescence ;
8 pubescent. Legs normal; teeth on hind femora varying from eight to eleven,
counting from base, 1 largest, 2 usually small and also usually the apical one or two.

d$.—Similar to the female, except that the pedicel is fuscous, also all the club and
preceding joint ; antennae as in the female, but the pedicel is smaller, the apical
funicle joint is as long as wide, and the club is somewhat shorter ; abdominal segments
5 and 6 comparatively longer, 7 shorter, the abdomen blunt at apex.

NORTH QUEENSLAND : Cairns district and Townsville.

Described from two females and one male received from Mr. G. F. Hill, and a large
number of females and one male collected by the author in January, March, May,
September, October and November. Mr. Hill’s specimens were bred from the
Curculionid beetle, Euthyrrhinus meditabundus. Of the author’s material, several

(2416) E2

68 ALAN P. DODD.

were bred from dead wood of the mango tree, and the remainder captured on dead or
dying timber frequented by the same weevil ; it was found easy to collect a series by
visiting the same tree for an extended period, one or more specimens being caught
daily.

The host record is of considerable interest, inasmuch as the other members of
the genus attack Lepidopterous and Dipterous larvae.

The diversity in habits of this species probably has some connection with the
several minor structural differences. The convex frons, narrow antennal scrobes,
concave occiput and produced scutum are peculiarities which might be given generic
value, but a study of a series of species would need to be entered upon. The author
has an unnamed species, collected in company with Chalcis euthyrrhint and probably
with similar habits, which shows the same characteristics.

There is little variation in colour ; the abdomen and posterior legs remain constant,
and the degree of variation in the first two pairs of legs and the antennae is not great.

Asymmetry in the femoral teeth is common; there may be nine teeth on one
femur and eleven on the other, or nine on one and ten on the other; the femora of
numerous specimens were examined.

Type. In the collection of the Queensland Museum, Brisbane.
Cotypes. In the collections of Mr. G. F. Hill and the author.

MOSQUITO NOTES.—II.

By F. W. Epwarps.
(Published by permission of the Trustees of the British Museum.)

Anopheles flaviceps, sp. nov.

Head with the integument rather light yellowish, especially when viewed from in
front. Erect scales mostly brown, a small area of white ones in the middle towards
the front, and in front of these a few long, narrow recumbent scales, not forming
a definite frontal tuft. Many light brown bristles between the eyes ; border bristles
dark. Antennae with the basal joint yellow ; in female with a few small scales on the
second joint ; male plumes light brown. Female palpi very slightly longer than the
proboscis, thin, with appressed scales ; narrow whitish rings at the tips of the first
three joints, just extending on to the bases of the succeeding joints ; tip of last joint
dark. Second joint slightly longer than the first, slightly shorter than the third
and fourth together ; fourth joint less than half as long as the third. [Male palpi
missing.| Pyvoboscis dark, except for the labella.

Thorax rather light yellowish, darker in the females (perhaps through discoloration).
Mesonotum somewhat shining, not darker at the sides, with light bristles, and in the
middle with rather numerous, very narrow hair-like scales. Prothoracic lobes rounded,
not mammillate, without scales. Prosternal hairs, five or six.

Abdomen light brown, with narrow dark bands in the male ; hairs pale ; no scales.
Male hypopygium: Basal spines, five or six, all rather strong, in a loose, irregular
cluster ; one distinctly distal to the rest and much longer, but not more slender.
Claspettes pointed ; a long and fairly strong apical hair, half as long again as the club,
which is normal in shape, but much more basally placed than the hair ; no accessory
hairs discernible. Tip of mesosome with about five rather narrow leaflets ; the longest
0-55 as long as the mesosome.

Legs dark ; extreme tips of femora and tibiae and extreme bases of tibiae pale ;
on the front and middle legs the tarsal articulations are also just perceptibly pale
under a strong lens. All femora slender, cylindrical.

Wings : Costa dark on basal two-fifths, in one wing of one specimen with a small
pale spot included near the base; apical three-fifths with four yellowish patches
alternating with three dark ones, of which the third is the smallest, the first being
distinctly larger than the adjacent light ones. A small dark spot at the wing-tip
opposite apex of upper fork-cell. First vein light with four dark patches, the first
being below the apical part of the dark basal two-fifths of the costa, the others below
the remaining dark costal spots. Remainder of wing without conspicuous markings,
but there are pale areas round the cross-veins, and the bases of the fork-cells, the base
of the second vein, the middle half of the third vein, and the extreme tips of the
second, third and fourth veins are also pale. Vein-scales linear, rather short and
not very dense.

ANGLO-EGYPTIAN SUDAN: Erkowit, 5.vi.1917, 1 3 (type), 1 2, also 1 9 from the
same place, 22.v.17 (dH. H. King).

Type in the British Museum, presented by the Imperial Bureau of Entomology.

Apparently most nearly allied to A. turkhudi, Liston, and A. hispaniola, Theo.
(which may be a local form of A. turkhudi), but differing in the paler colour of the
integument, especially of the head, almost obsolete frontal tuft, more numerous
prosternal hairs, more shining mesonotum, and somewhat less conspicuous wing
markings ; also in the male hypopygium in the more pointed claspettes, which have
no accessory hair and a more basally situated club, and in the stronger detached
basal spine.

70 F. W. EDWARDS.

Anopheles immaculatus, Theobald.

Swellengrebel and Swellengrebel de Graaf have recently suggested (Bull. Ent? Res.,
xi, p. 78, 1920) that this may be an albinoid form of A. vagus, Dénitz. An examination
of the hypopygium of a male from Java presented to the British Museum by
Dr. Swellengrebel supports this possibility, since no tangible differences could be
discovered between the hypopygia of the two forms.

Anopheles leucosphyrus, Donitz.
Two rather well-marked forms of this species occur :—

(1) The typical form. Female palpi with the white rings distinct, the last joint
white on at least its apical half. Proboscis of normal length, little, if any, longer than
the palpi. Dark markings of wings less extensive ; the spots on the first longitudinal
vein more broken up.

(2) Var. hackert, nov. Female palpi with the white rings very narrow, the last
joint white only at the extreme tip. Proboscis unusually long, longer than the palpi
by almost, or quite, or even more than, the length of the last two palpal joints. Dark
markings of wings more extensive ;_ the spots on the first longitudinal vein more fused.

The var. hackeri has recently been noted by Dr. H. P. Hacker (Fed. Malay States,
Malaria Bureau Reports, ii, p. 33, 1921). A specimen collected by him and presented
to the British Museum is designated as the type ; the Museum collection also contains
a number of others from the Malay States, from Dr. Leicester’s collection. The
distinction in colour between the palpi of the two forms is quite sharp, and in length
also it is very striking, but variable. Most specimens of the typical form have the
palpi as described, almost, or quite, as long as the proboscis. However, among a small
number from Borneo collected by Dr. Roper there is one which has an elongate
proboscis like that of the dark form. Probably, therefore, the two forms are not
specifically separable.

Anopheles punctulatus, Dénitz.

This species has recently been discussed by Swellengrebel and Swellengrebel de
Graaf (Bull. Ent. Res., xi, p. 89, 1920), who concluded that it was not specifically
distinct from A. tesselatus, Theo., and at the same time described a very similar
form under the name Nyssorhynchus annulipes var. moluccensis. Finding that the
description of moluccensis agreed rather closely with my conception of punctulatus,
I wrote to Dr. Swellengrebel, suggesting that the two were the same, and received the
following reply :—

“As to Dénitz’s punctulata, there can be no doubt, judging from the published
photograph, that its proboscis is white on the apical half. Moreover, the black ring
near the apex of the second palpal joint is very narrow. This induced me to separate
moluccensis from it, and I still think that the specimen from which the photograph
was taken (probably the type) is a tesselata. Asa general rule I believe it is well, from
a practical point of view, to separate these allied forms, if, at least, a separation is
practicable ; some of them may prove to be good carriers and others not (vossiz and
ludlowt !).”’

After a close study of the literature and the British Museum collections, somewhat
different conclusions seem to be indicated from those arrived at by Dr. Swellengrebel.

All the specimens in the Museum series of A. punctulatus prove to be practically
identical with moluccensis in palpal markings, and most of them also agree in having
the proboscis entirely black. One or two, however, including a specimen determined
by Dénitz and coming from his type locality (Stephansort), show a pale area on the
underside of the proboscis towards the tip. The colour of the proboscis cannot,
therefore, always be used to separate moluccensis from tesselatus. I cannot agree

MOSQUITO NOTES—II. 71

with Swellengrebel in placing moluccensis as a variety of annulipes; the latter,
apart from its much larger size, shows some quite good differences in the wing-
markings.

The three forms under consideration appear to be separable as follows (in the
female sex) :—

(1) A. punctulatus, Dénitz, typical form. Proboscis pale on the apical half, at least
on the underside. Second joint of palpi about equal in length to the third and fourth
together ; its basal half black, and a narrow dark ring close to the tip of the white
apical half. New Guinea.

There is no specimen exactly answering to this description in the British Museum,
and Swellengrebel states that he has not met with it. Probably, therefore, Dénitz
may have described and figured a somewhat aberrant specimen of the following.

(2) A. punctulatus var. moluccensis (Swellengrebel). Proboscis entirely black, or
with a rather small pale area on the underside towards the tip. Second joint of palpi
markedly longer than the third and fourth together ; its basal three-fifths or more
black, and a broader black ring towards the tip. New Guinea; Moluccas.

This is A. punctulatus in the sense in which I have previously understood it.

(3) A. punctulatus var. tesselatus, Theobald. Proboscis pale on the apical half
(above and below). Second joint of palpi markedly longer than the third and fourth
together ; its basal half black, its apical half entirely white. Oriental Region ; also
reported by Swellengrebel from Ceram, but this may prove to be the true punctulatus.

Anopheles amictus, sp. nov.

Differs from A. annulipes, Walker, as follows :—Proboscis entirely dark-scaled in
both sexes. Antennae of the female shorter and stouter, especially the last few
joints, which are neither thinner nor longer than those immediately preceding, the last
ten joints all being of about the same length and thickness. Abdomen rather densely
clothed with broad, flat scales on all segments except the first ; on the dorsal surface
the scales are mostly yellowish, with a patch of dark ones in the middle of segments
4-6 ; on the ventral surface the scales are mostly whitish and less numerous towards
the base. White spots on the legs larger, the posterior surface of the front tibiae
being almost entirely white. First longitudinal vein with 14-18 small black spots ;
no long ones.

QUEENSLAND : Townsville (G. F. Hill), type 2 and one 6, presented by the
Imperial Bureau of Entomology.

The differences between this and A. annulipes seem to be too great to fall within
the limits of specific variation, especially as regards the abdomen and antennae ;
nevertheless, the two are certainly very closely allied, and some specimens show
intergradation in some respects. A female from Townsville, 2.ii.1903 (F. P. Dodd),
agrees with the above definition, and also shows a further difference from A. annulipes
in that the second, third and fourth joints of the hind tarsi have distinct whitish
rings at the base as well as at the tip. A female from Port Darwin agrees with this.
A female from Townsville (’. H. Taylor), and another from Cardington, Queensland
(F’. H. Taylor), have the scaly abdomen of A. amictus, but the antennae are somewhat
more slender apically, the proboscis is pale on the apical half, the integument of the
mesonotum is ochreous, and the first vein has long black spots. The last two specimens
may possibly be A. mastersi, Skuse, but regarding both A. masterst and A. musivus
Skuse states that the /ast abdominal segment bears scales, implying that the others
do not. Besides the above-mentioned forms, examples of normal A. annulipes have
been taken at Townsville.

TBA F. W. EDWARDS.

Megarhinus (Toxorhynchites) kempi, sp. nov.

3. Head blue-scaled (? rubbed). Proboscis purple, the thin portion with a greenish
tinge. Palpi rather longer than the proboscis, slender ; first three joints about equal
in length, mainly yellow-scaled, some purple scales, chiefly on upper surface, towards
bases of first and second, and at apices of second and third. Fourth joint dark,
acuminate, nearly as long as second and third together. Clypeus and basal antennal
joints bluish-grey-dusted, shaft of antennae yellow ; first flagellar joint witha few dark
scales. Thorax: Prothoracic lobes clothed with deep blue scales, a few white ones
beneath. Pro-epimera and pleurae with silvery-white scales. Mesonotum black,
scales mostly metallic green, with some purple ones intermixed (very much rubbed).
Abdomen without apical lateral scale tufts ; dorsal surface mostly metallic purplish
blue; first tergite green; remaining tergites each with a narrow basal band of
bluish green. Venter mostly yellow ; no median purple line ; eighth sternite purple.
Legs: Femora purple above and in front, yellow beneath. Tibiae purple. Tarsi
purple, the first joint on all the legs with a rather narrow and ill-defined whitish ring
near the base. First hind tarsal joint with rather long and dense bristles beneath,
except at the base. Front and middle claws unequal, the larger with a strong median
tooth, the small simple ;_ hind claws equal and simple. Wings as in M. leicester1.
Hypopygium: Ninth tergite rather narrow, the bare middle part slightly emarginate
apically, fully twice as wide as long ; side portions each with 10-12 long hairs. Ninth .
sternite bare. Side-pieces tapering, nearly three times as long as their basal width ;
basal lobes triangular, apically with two long and strong bristles. Claspers slightly
shorter than the side-pieces, of even width throughout, terminal spine long, strong and
pointed ; about five fine hairs in a row before the tip. Tenth sternites long and rather
slender, their tips not conspicuously enlarged or blackened. Lower bridge of mesosome
very narrow and nearly basal ; paired processes of mesosome rather slender, distinctly
but irregularly serrate above (sternally).

2. Palpi purple, not longer than the head. Head with mixed colours, mostly
blue above and pale round the eyes. Prothoracic lobes purple. Mesonotal scales
bright metallic green. White rings on the first tarsal segments more distinct than in
the male; the tarsi have also additional white markings as follows :—cn the front
and middle legs, a small area at the base of the second joint on the underside ; on the
front legs, the inner and lower side of the fourth joint, except its tip, and the underside
of the fifth joint ; on the middle legs, practically the whole of the fourth and fifth
joints, except the tip of the latter on the upper side. (The last four joints of the hind
tarsi are missing.)

Inp1A: Talewadi, nr. Castle Rock, N. Kanara district, 3-10.x.1916 (S. Kemp),
1919. Type in the Indian Museum.

Closely resembles M. leicesteri, Theo., which differs chiefly in having the whole of
the last four joints of the middle tarsi of the female white ; the ninth tergite of the
male is broader, and the paired anterior processes of the mesosome are not serrate.

Megarhinus (Toxorhynchites) klossi, sp. nov.

3. Differs from M. kempi as follows :—First joint of palpi a little shorter, third a
little longer than second. Scales of prothoracic lobes all purplish blue; those of
pro-epimera coppery above ; of mesonotum duller, purplish and bronzy, mixed with
green ; of scutellum coppery. Sides of first abdominal tergite whitish. Ninth tergite
broader ; tenth sternites more enlarged apically ; lobes of mesosome smooth above ;
clasper with hair-like terminal spine. First hind tarsal joint without dense bristles
beneath. From T. gravelyi the differences are :—Distinct basal bands are present on
the abdominal tergites ; ninth tergite is less emarginate ; venter is without median
purple stripe, etc.

Frep. Matay States: Kedah Peak (Gunong Jerai), 3,200 ft., xi—-xii.1915
(C. Boden Kloss), 1 3. Type in the British Museum.

MOSQUITO NOTES—II. is

Megarhinus (Toxorhynchites) gravelyi, sp. nov.

3. Head mostly dark-scaled (colour varying with the light), a pale rim round the
eyes. Proboscis purple, more greenish on the thin apical portion. Palpi slender,
slightly longer than proboscis ; first joint distinctly shorter than the second or third,
which are about equal in length, and together about as long as the fourth ; the palpal
scales are purple, on the underside of the first three joints, except towards the tip of
the third and the base and extreme tip of the second, mostly golden. Second antennal
joint with some golden scales. Thorax: Prothoracic lobes and pro-epimera coppery-
scaled above, silvery white below ; pleurae silvery-white-scaled ; mesonotal scales
bright metallic green ; scutellar scales coppery, a few whitish ones at the sides. Abdo-
men: First tergite bluish green in the middle, shining creamy white at the sides ;
remaining tergites deep blue, with basal lateral creamy spots. A few yellow hairs at
the sides of tergites 6 and 7, not forming definite tufts. Venter golden, with a narrow
median purple line; eighth sternite purple. Legs purple-scaled; femora golden
beneath and towards the base ; first joint of all tarsi with a narrow indistinct pale
ring near the base (joints 3-5 of hind tarsi denuded). First hind tarsal joint without
dense bristles beneath. Wings with the cross-veins nearly in a line, the m-cu cross-
vein somewhat oblique outwardly. Hypopvgium : Ninth tergite strongly emarginate
apically, narrow.in the middle, with well-developed hairy lobes, each bearing 10-15
hairs. Ninth sternite bare. Side-pieces tapering, hardly more than twice as long as
their width at the base. Basal lobes with three strong bristles, one weaker than the
others. Tenth sternites stout, blackened and somewhat enlarged apically. Claspers
as long as the side-pieces, slightly tapering at the tip, terminal spine long, strong,
pointed. Mesosome with the lower bridge very narrow, basal; lobes rather slender,
smooth above.

Inp1IA: Pashok, Darjiling district, 2,000ft., E. Himalayas, 26.v.—14.vi.1916
(Ff. H. Gravely), 1 3.

Type in the Indian Museum.

Resembles 7. metallicus, Lin., differing in the darker upper side of the palpi, coppery
rather than purple scales on upper part of prothoracic lobes and pro-epimera ; presence
of pale rings on first tarsal joints, and more emarginate ninth tergite.

Opifex fuscus, Hutton.

This species was originally described from New Zealand by Hutton (Trans. N.Z.
Inst., xxxiv, p. 188, 1902) as a Tipulid, and its true position has not till now been recog-
nised. Recently a number of specimens have been presented to the British
Museum by Mr. G. V. Hudson, taken on the ocean beach at Wellington, N.Z.

The insect is a Culicine mosquito, presenting many remarkable features ; though its
peculiarities seem to be mainly connected with sex, it certainly cannot be placed in
any previously known genus of Culicidae, and it is difficult to place it precisely in
relation to the other genera of the Curtcinr. It seems to be nearer to Aédes than
to Culex. The following are the most striking features :—

Male.—Antennae rather stout, not plumose, the terminal joints not lengthened ;
the first joint conspicuously hairy ; the third, fourth and fifth joints a little shorter
than those which follow, and each bearing a strong spine at the base on the upper side,
the spine on the fifth joint very long. Proboscis and palpi strongly curved ; the palpi
two-thirds as long as the proboscis, the last joint forming a club. The head has no
upright scales, these being replaced by hairs. Eyes well separated. Abdomen in
several specimens curled under the thorax. Hypopygium with the side-pieces
simple, broad at the base, pointed at the tips, a membranous strip along the inside
from base to tip ; clasper subapical, short, with strong terminal claw and sub-basal
projection ; anal segment well developed, tenth tergites hairy, tenth sternites each
ending in a single strong sharp point ; aedeagus of simple structure, resembling that

74 F. W. EDWARDS.

of Ochlerotatus. Femora and tibiae stout, especially on the front legs, where the tibiae
are swollen and very short, not more than two-thirds as long as the femora. _ Fifth
joint of front tarsi very small, not longer than broad, but the claws enormous, longer
than the last three tarsal joints together, equal, simple, divaricate. Claws of middle
and hind legs moderately large, equal and simple. Wings as in Aédes.

Female.—Antennae and front legs not specially modified; claws all simple.
Palpi about a quarter as long as the proboscis. Abdomen rather blunt; cerci
rather short and broad ; eighth sternite very large.

The peculiar antennae and front legs of the male must have some special biological
significance, and it is to be hoped that New Zealand collectors will be able shortly
to work out the life-history of the insect.*

Leicesteria annulipalpis (Theobald).

In my paper on the genus Leicesteria (Bull. Ent. Res., iv, pp. 255-263, 1914) I
expressed doubt as to the correct location of L. annulipalpis. Recently, however, a
female has been received from Dr. N. H. Swellengrebel, taken at Mandailing, Sumatra,
which shows that the species really is a true Leicesteria. This Sumatran specimen
agrees with Theobald’s description, except that the claws show a slight swelling
towards the middle, which evidently represents a tooth.

Aédes (Stegomyia) dendrophila, sp. nov.

Closely related to S. fraseri, Edw., differing almost solely as follows :—Middle:
femora without a white spot in the middle in front. Hypopygium of male with the
side-pieces shorter, less than twice as long as their breadth at the base ; basal lobes
larger, more densely hairy; claspettes entirely unrepresented ; claspers shorter,
straighter, and less attenuated apically.

GoLtp Coast: Nsawam, 16.11.1920 and 14.iv.1920, 2 3, 3 9, reared from larvae
in hole in cotton tree ; Oblogo, 17.iv.1920, 3 gf (including type), 2 2, from tree-hole ;
Aburi, 6.vi.1920, 5 3,2 9, from banana (Dr. A.Ingram). SIERRA LEONE: Freetown,
ix.1914, 1 Q from tree-hole (Dr. G. Butler, presented by A. Bacot) ; previously
recorded as S. fraser.

The absence of a white spot on the mid femora would place this species with
S. pseudonigeria in my key (Bull. Ent. Res., iii, p. 8, 1912). The new species differs
from S. pseudonigeria as follows :—All tibiae with small whitish spots at the extreme
base, not removed from the base, beneath ; mid femora all black on posterior surface,
except for the apex ; second joint of mid tarsi all yellowish white, except beneath
towards tip (in S. psewdonigeria white at the base only) ; last two joints of hind tarsi
with some dark scales beneath ; segments 2-5 of abdomen with narrower whitish
basal bands or none.

Aédes (Ochlerotatus) bancroftianus, sp. nov.

Head clothed almost entirely with broad, flat scales, a rim of narrow ones round the
eyes ; the flat scales varying in colour from dark brownish to cream-coloured, generally
paler towards the nape; the narrow scales ochreous; ocular bristles pale. Torus
ochreous, darker on the inner side. Proboscis dark-scaled and slender throughout,
one-third longer than the front femora. Palpi dark-scaled ; in the female about one-
sixth as long, in the male of exactly the same length as the proboscis ; last two joints
in the male slightly swollen, with well-developed hair-tufts, the terminal a little shorter
than the penultimate. Thorax brown, mesonotum clothed in the type with moderately

* Since writing this I learn that Mr. D. Miller, Government Entomologist in New Zealand, has
an account of the life history of this insect in course of publication. Larvae and pupae of the
species have been received from Mr. G. V. Hudson; they show some resemblances to Armigeres.

MOSQUITO NOTES—II. 75

dark brown narrow scales ; ochreous narrow scales round the front margin, round the
ante-scutellar space, and in two lines extending from the scutellum for nearly half
the length of the mesonotum ; in other specimens the dark scales are lighter and the
pale scales are more numerous, forming an indistinct pale median transverse band.
Scutellum with narrow pale ochreous scales. Prothoracic lobes with broadish curved
scales ; pleurae with broad flat white ones. Abdomen ochreous, with dark brown
scales dorsally ; tergites 2-4 with complete basal creamy bands, broadening some-
what laterally, 5 and 6 with lateral basal creamy spots. Venter pale-scaled, sternites
4-6 with dark apical bands. In the female the seventh segment is remarkably
small, narrow, and partly retracted ; the cerci are long and narrow, nearly three times
as long as broad. Male hypopygium: Side-pieces nearly cylindrical, almost four
times as long as their width at the base, without lobes, but with an aggregation of
hairs into a small dense patch at the base of the lower flap. Claspettes entirely
unrepresented. Claspers long, the outer third rather suddenly narrowed and curved
inwards ; terminal spine long, slender, almost straight. Lobes of ninth tergite small,
with a few short bristles. Mesosome rather short and broad, simple. Anal segment
normal. Legs entirely dark-scaled, except for the undersides of the front and middle
femora, and the greater part of the hind femora, except the tip and a line along the
upper side of the outer half. Claw-formula :— $§2:1.1:1.0:0; 21:1.1:1.0-0.
Wings entirely brown-scaled ; scales of the lateral series long, linear, those of the
median series also rather long. Cell R, as long as its stem in the male, longer in the
female, its base slightly proximal to that of cell M,. Halteres ochreous, knob
somewhat darker. Wing-length 3-5-4 mm.

QUEENSLAND: Eidsvold (Dr. T. L. Bancroft). Type female, 12 other females,
and 3 males presented to the British Museum by the Imperial Bureau of Entomology ;
received through Mr. G. F. Hill.

The only near ally of this species is O. multiplex, Theo., which differs in thoracic
ornamentation, in the larger seventh segment of the female abdomen, and in the
straighter and nearly cylindrical male claspers.

Aédes (Ochlerotatus) ashworthi, sp. nov.

Head clothed at the sides with broad flat whitish scales ; in the middle of vertex
with loosely applied narrower flat straight scales; on the nape with quite narrow
curved scales; bristles black. Proboscis dark-scaled, slender, nearly one-half longer
than the front femora. Palpi dark-scaled ; in the female nearly one-fourth as long as
the proboscis, the second joint swollen and longer than the first ; in the male about
four-fifths as long as the proboscis, long joint almost divided in the middle, equal in
length to the last two joints together, tip of long joint and whole of penultimate joint
with long dark hair, terminal joint very broad, slightly shorter than penultimate,
nearly bare. Thorax : Integument of mesonotum dark brown, with large pale humeral
patches, and traces of three blackish longitudinal lines ; mesonotal scales all narrow,
mostly ochreous; with some black ones intermixed. Pleurae with some flat, dull
whitish scales. Abdomen brown, a broad band of dull grey scales at the base of each
segment. Seventh segment of female large; eighth sternite also rather large and
prominent ; cerci very small, not longer than broad. Male hypopygium: Lobes of
ninth tergite clothed with short hairs. Side-pieces about 3-5 times as long as broad,
the lower flap without lobes, but bearing on its edge a row of peculiar bristles which are
flattened towards the tips, almost club-shaped, these bristles extending for more than
two-thirds of the distance from the base of the side-piece. Claspette represented by
a small basal lobe of the upper flap of the side-piece, this lobe truncating apically and
bearing about a dozen rather long twisted scales or flattened bristles. Claspers long,
nearly cylindrical, gradually narrowed and somewhat curved on the apical half, with
a moderately long terminal spine. Mesosome rather short and broad, simple. Legs
dark, except for the whitish undersides of the femora and the narrowly ochreous tips

76 F. W. EDWARDS.

of the femora and tibiae. Claw formula? @2:1.2:1.121; (90) 171) 1-48 |
Wings entirely brown-scaled ; scales of the lateral series long, linear; those of the
median series also rather long. Cell R, over twice as long as its stem in the female,

its base slightly proximal to that of cell M,. Wing-length 4-4-5 mm.

Larva: Head rather broad for this group; clypeal hairs all simple. Antennae
moderately long, slightly curved, almost cylindrical and nearly bare ; a single hair |
just beyond the middle. Comb of eighth abdominal segment consisting of a triangular |
patch of 50 or more pointed scales. Siphon about three times as long as its greatest

breadth, slightly contracted at base and tip, not very strongly chitinised. Pecten

short, consisting of about 12 (fewer in the younger larvae) sharp-pointed, simple |
teeth, placed so close together that their bases touch ; the first of these teeth is situated _
at about one-fifth of the length of the siphon, but in some specimens the actual pecten _
is preceded by one or two small, widely-spaced, simple bristles. Hair-tuft of about |
eight hairs, situated a little beyond middle of siphon. Tracheal tubes very large, |
strongly chitinised, occupying almost the whole of the middle part of the siphon, but |
strongly and abruptly contracted before the base of the latter. Anal segment with |

a lightly chitinised saddle; gills very small and globular. Brush well-developed,
with 10-12 elements.

West Australia: Yallingup, ix.1914 (Dr. J. H. Ashworth). Type and one |

other male, one female, and a dozen larvae presented to the British Museum by the
Imperial Bureau of Entomology in 1915.

This species is nearly allied to O. crucians, Walker (tasmaniensis, Strickland), and
O. (Caenocephalus) concolor, Taylor, differing from both chiefly in the details of the
hypopygium. O. crucians has the clasper very much swollen in the middle, its
terminal spine shorter and stouter, and the flattened bristles on the margin of the side-
pieces much less numerous ; the terminal joint of the male palpiis much more slender.
Probably the specimens of O. concolor from Tasmania recorded by Taylor are really
O. crucians ; the true O. concolor from New South Wales (which was wrongly described
as having simple female claws) resembles O. crucians in its hypopygium, but the
claspers are less swollen, the scales on the claspette lobes are less numerous, and the
last joint of the male palpi is swollen, as in O. ashworthi. Doubtless the three forms
are geographical representatives of the same type.

Aédes (?Skusea) funerea, Theobald.

A male and female, apparently of this species, have been received from Dr. Swellen-
grebel, from Amboina. No Australian male is available for comparison, and the
identification is therefore not absolutely certain, though probably correct. -The male
hypopygium of the Amboina specimen does not show the least resemblance to
S. pembaensis, but, on the other hand, resembles that of A édes panayensis, Ludlow ;
from this it differs in the absence of a long process at the base of the side-piece, and
in the presence of a short spine instead of a thick projection at the tip of the side-
piece. Probably the species would be better placed in the subgenus Aédes than in
Skusea, but much more knowledge of the biology of these forms is necessary before
their classification can be regarded as satisfactory.

Aédes (? Skusea) funerea var. ornata, Theobald.

Lepidotomyia lineata, Taylor, Trans. Ent. Soc., 1914, p. 191 (1914).

A male and female have been received from Dr. Swellengrebel from Ceram. The
male hypopygium is identical with that of S. funerea from Amboina, and the form is
therefore probably correctly regarded as merely a variety of S. funerea, in spite of the
striking difference in thoracic ornamentation. Two female paratypes of L. lineata,
sent by Mr. G. F. Hill, agree with the female from Ceram.

MOSQUITO NOTES—II. 77

Aédes (Skusea) punctipes, sp. nov.

Q. Head entirely clothed with rather close-lying, flat blunt-ended scales: on the
upper surface three black patches alternate with four white ones, black scales occurring
again low down at the sides. Eyes separated by a rather narrow white-scaled line.
Proboscis dark-scaled, not swollen at the tip, equal in length to the front femora.
Palpi dark-scaled, rather more than a quarter as long as the proboscis. Antennae
missing. Yhorax : Prothoracic lobes and pro-epimera with rather broad white scales ;
mesonotum with narrow curved white scales round the front margin, narrow light
bronzy-brown scales rather densely covering the rest of the surface ; these scales are
somewhat broader and denser on the posterior portion of the mesonotum, while on the
space in front of the scutellum, which is normally bare, as well as on the scutellum
itself, are broad flat blackish-brown scales ; a small patch of similar scales above the
root of each wing. Postnotum bare. Pleurae with patches of flat white scales. Meso-
notal bristles mostly denuded, probably rather long and dense. Abdomen brown-
scaled above, except the first tergite, which has creamy scales; tergites 2-7 have
lateral white spots, which are not quite basal in position and extend a short way on to
the dorsal surface. Venter mostly pale, apical sternites dark-scaled apically. The
eighth segment and cerci are not visible externally ; sixth and seventh sternites
prominent in side view. Legs mostly brown-scaled ; femora lighter beneath, and with
a distinct whitish preapical ring; extreme tips of femora and tibiae also whitish.
Front tibiae with three, middle and hind tibiae with four small, but distinct, whitish
spots on the anterior surface. First joint of all tarsi with a narrow white ring at the
base ; junction of first and second joints with a small white dorsal spot ; first mid-
tarsal joint also with a median dorsal white spot, which on the hind legs becomes a
complete narrow white ring; second hind tarsal joint narrowly white at the tip.
Tibial bristles short, pale. Mid and hind tibiae equal in length. First tarsal joint
on front legs about half as long as the tibia, and distinctly shorter than the remaining
joints together ; on hind legs about two-thirds as long as the tibia. Claws simple.
Wings brown-scaled, except for the base of the fifth longitudinal vein, which is white.
Scales all rather long and broad, mostly obliquely truncate at the tips; only a few
longer and narrower ones towards the tips of the veins and along the lower margin of
the upper branch of the fifth vein. Wing-fringe very long. Upper fork-cell fully
twice as long as its stem, and with its base nearer the base of the wing than that
of the lower.

Upper Burma: Maymyo, xi-xii.1913 (Major Bennett), 1 9.

Type in the Indian Museum.

A very distinct species, easily recognised by the leg markings and thoracic scaling.
The structure of the abdomen and claws shows that it is quite closely related to
Stegomyia periskeleta, Giles (=Ochlerotatus annulifemur, Edw.), and S. microptera,
Giles. Both these species have male palpi of the Ochlerotatus rather than of the
Stegomyia type, and, on the other hand, show some relation to Armugeres in the
structure of the mesosome of the aedeagus. The type species of Skusea (S. pembaensis,
Theo.), though with very dissimilar hypopygium, agrees with these Oriental species
in general appearance and in the structure of the male palpi and female abdomen and
claws, and all four are probably best relegated to the same genus or subgenus of the
Aédes group. The new species, like its ally S. microptera, may be expected to be a
tree-breeder. It is in such habitats that the species showing the most striking varia-
tions from the normal type of Aédine structure are found, sometimes, as in the present
instance, indicating connections with other groups.

Culex crinicauda, nom. nov.
Culex parvus, Taylor, Bull. N. Terr. Austral.; la, p. 27 (1912) ; nec Culex parvus,
Macquart.
Although my suggestion that this might be synonymous with C. vishunt, Theo.,
was adopted by Taylor, such is, nevertheless, not the case. The hypopygium of a

78 F. W. EDWARDS.

male determined by Taylor and sent by Mr. G. F. Hill is very different from that of
C. vishnui, and shows some peculiar characters. The clasper is unusually broad, almost
straight, and has round its base a rather dense tuft of hairs. The lobe of the side-piece
bears the usual leaf and filaments, but the filament adjacent to the leaf is remarkably
long, flattened, and backwardly (caudally) directed.

Culex taylori, nom. nov.

Leucomyia annulivostris, Taylor, Trans. Ent. Soc. 1913, p. 696 (1914) ; nec Culex
annultrostris, Skuse.

A male paratype has been received from Mr. G. F. Hill. The species appears to be
a distinct one, allied, as Taylor stated, to C. sinensis, Theo., but differing in having
the pale bands of the abdomen confined to the bases of most of the segments, and
without pale spotting on the tibiae. From C. sitiens it differs in the much broader
pale apex to the last palpal joint.

Culex basicinctus, nom. nov.*

Leucomyia annulata, Taylor, Trans. Ent. Soc. 1913, p. 695 (1914); nec Culex
annulatus, Schrank.

Two males and three females have been received from Mr. G. F. Hill, collected by
him at Townsville. They were sent as Leucomyia annulirostris, but agree with Taylor’s
description of L. annulata, and are almost certainly that species. C. basicinctus
seems nearly allied to C. whttmoret, Giles, and, like that species, has the pale rings of
the tarsi confined to the bases of the joints, and the pale bands of the abdomen
angularly produced in the middle ; it differs obviously in many points, e.g., its larger
size, duller thoracic colouring, and broader white tip to the last joint of the male palpi.
A peculiar character, not mentioned by Taylor, is that the male palpi have two black
bristles at the extreme tip, which are conspicuous against the adjacent white hairs.

Culex ventrilloni, Edw.
Culex ventrillont, Edwards, Bull. Ent. Res. xi, p. 135 (Sept. 1920).
Culex albigenu, Enderlein, Wien. Ent. Zeitschr. xxxviii, p. 50 (Nov. 1920).

A male of C. albigenu lent me by Dr. Enderlein enables me to state the above
synonymy. Somewhat unexpectedly, the species proves to be a true Culex, in Dyar’s
most restricted sense; the hypopygium has almost the same structure as in the
African C. stmpsoni, Theo., and C. andersoni, Edw.

Culex quasigelidus, Theo.
Culex auritaenia, Enderlein, Wien. Ent. Zeitschr. xxxviii, p. 49 (Nov. 1920).

The above synonymy, evident from the description, is confirmed by the examination
of specimens sent by Dr. Enderlein.

Culex (Lophoceratomyia) jenseni (Meij.).

Cyathomyia jensent, de Meijere, Ann. Jard. Bot. Buitenzorg, (2) iii, p. 922 (1910).

Lophoceratomyia curtipalpis, Edwards, Bull. Ent. Res. v, p. 127 (1914).

The above synonymy is proved by comparison of the type of L. curtipalpis with a
paratype of C. jenseni, presented to the British Museum by Professor de Meijere.
The species is wrongly placed in my table of the species of this subgenus (Bull. Ent.

Res. vii, p. 227, 1917) ; it should come under heading 9, differing from the other species
there included in the shorter male palpi.

* (There does not appear to be any reasonable ground for proposing this new name. Schrank’s
species was transferred to Theobaldia long before Leucomyia annulata was described, and there
can be no possibility of confusion between them.—Ed.]

MOSQUITO NOTES—II. 79

My previous use of the name Cyathomyia in the sense of Pvotomelanoconion,
‘Theobald, proves therefore to be erroneous ; if the group is retained as a distinct
subgenus of Culex, Theobald’s name will have to be revived, Cyathomyia falling as a
synonym of Lophoceratomyia. However, C. brevipalpis, Giles, and P. fuscum, Theo.,
have a hypopygium constructed much as in Neoculex, Dyar, and should probably be
referred to that subgenus.

Rachisoura filipes (Walker).
Culex filipes, Walker, Proc. Linn. Soc. v, p. 229 (1861).
Rachisoura sylvestris, Theobald, Mon. Cul. v, p. 208 (1910).
Stegomyia hilli, Taylor, Proc. Linn. Soc. N.S.W. xxxix, p. 456 (1914).
Mimeteomyia hilli, Taylor, Proc. Linn. Soc. N.S.W. xli, p. 566 (1916).

The above synonymy appears to be proved by a comparison of Walker’s and
‘Theobald’s types with a specimen of S. fall: from Stapleton, Northern Territory,
sent by Mr. G. F. Hill. The genera Rachisoura and Mimeteomyia are probably not
distinguishable ; they differ from Rachionotomyza in the shorter and stouter proboscis,
which is little, if any, longer than the front femora.

Rachionotomyia aenea, sp. nov. 2.

Head dark-scaled, with a moderately broad pale band in front, which is blue or
whitish according to the direction of the hight. Palpi and proboscis purplish-scaled,
the palpi exceeding the clypeus by about twice the length of the latter. Antennal
torus orange. Thorax with the integument orange-brown, darker brown on the
mesonotum. Prothoracic lobes with black bristles and narrow, almost hair-like,
dark brown scales. Mesonotum with narrow dark brown scales, except on, and just
in front of, the scutellum and between the scutellum and the wing-bases, where the
scales are broad, flat and bronzy-green in colour. Pleurae with a large silver-scaled
dark brown patch. Abdomen purplish-black above, golden beneath, the tergites with
lateral subapical bluish-silvery patches. Legs purplish-scaled ; middle femora in
front with a silvery streak on the basal half and a silvery subapical spot ; front and
hind femora unmarked. Wings with dark brown scales; those on the fork-cells
short and rather broad. Upper fork-cell slightly longer than its stalk, its base™
distinctly distal to that of the lower. Cross-veins separated by more than the length
of the posterior. Wing-length 3 mm.

Matay STATES: Edges of stream, Ampang jungle, Kuala Lumpur, 21.v.1904
nor Ge Fa Letcester), | 9.

Allied most nearly to R. similis, Leicester, differing in the unspotted front and hind
femora, and in some other points.

Rachionotomyia purpurata, sp. nov. 2°.

Nearly allied to R. bimaculipes (Theobald), differing as follows :—Integument of
mesonotum orange, almost dull (instead of shining blackish), clothed with narrow
greenish scales mixed with some black ones ; pro-epimera with small flat black scales
(bare, perhaps rubbed, in R. bimaculipes) ; dark scales of abdomen with strong purple
reflections (instead of dull black).

Fit Is.: Suva, 10.iv.1911, 3 2 reared from larvae (Dr. P. H. Bahr, pres. by
Lt.-Col. A. Alcock).

80. .F. W. EDWARDS.

Rachionotomyia quasiornata (Taylor).
Stegomyia quasiornata, Taylor, Proc. Linn. Soc. N.S.W. xl, p. 177 (1915).

This is very similar to R. bimaculipes and R. purpurata, but differs from both in
having narrow instead of broad and flat scales on the prothoracic lobes. A female
has recently been received through the Imperial Bureau of Entomology, named by
Mr. G. F. Hill after comparison with Taylor’s type; it shows the long proboscis
characteristic of the genus. In all these three Australasian species the palpi are
shorter than in the Oriental forms, exceeding the clypeus by hardly more than the
length of the latter.

81
BIOLOGICAL STUDIES OF APHIS RUMICIS, LINN.*

By J. Davipson, D.Sc.

The Entomological Department, Institute of Plant Pathology
Rothamsted Experimental Station, Harpenden.

This is the first of a series of papers based on results of breeding experiments and
on observations in the field. It is hoped that a full investigation of the biology of this
species will be of value in elucidating the many difficult biological problems of the
APHIDIDAE.

The preliminary paper published by the author (1914) was to have been followed by
further investigations, but owing to the outbreak of the European War there has
been an unavoidable delay. However, experiments are now being continued, and the
results of the researches will be published in parts from time to time.

As there is no complete description of Aphis rumicts in the literature, it is thought
very desirable that a detailed illustrated description of all forms of this species should
be given.

The following abbreviations are used in the text :—a.v. 9 = apterous viviparous

female ; w.v. 2 = winged viviparous female ; Ist v. gen. = Ist, 2nd, etc., viviparous
generation.

I. Description of Aphis rumicis, Linn.
1. FUNDATRIX (fig. 1). Average size, 1:8 mm. by 1-4 mm.f

Body oval to elongate, broadly rounded posteriorly, shorter and relatively stouter
than the succeeding a.v. 2 2; colour black to dark green; hairs scattered over
the body.

Head: A few scattered hairs on dorsal surface. Eyes black ; small tubercle-like
accessory eyes{ on posterior margin. Antennae§ about two-thirds length of body ;
black to dark brown, paler about the middle ; five segments ; seg. 3 the longest ;
1 slightly broader than long; 2 slightly longer than broad, subequal in length ;
4‘shorter than 5; 5 almost as long as 3; a single subapical sensorium on seg. 4 and
a compound sensorium on seg. 5; a few short hairs on each segment. Rostrum
normal, with a few hairs on each segment.

Thorax: A pair of prominent lateral tubercles on prothorax. Legs black, with
tibiae and proximal portions of femora paler ; segments bearing many short hairs,
especially the tibiae.

Abdomen with two prominent lateral tubercles on each side. Cornicles black to dark
brown, imbricated, tubular, tapering very slightly distally, shorter than in succeeding
a.v. 2 2; about one and one-third times thelength of cauda as seen from the dorsum.
Cauda short, bluntly rounded, black on distal portion ; bearing several long curved
hairs. Anal plate black, roughly quadrangular as seen from venter, bearing a
number of short hairs, anterior margin more or less straight. Genital plate black,
somewhat crescentic in shape, with outer margins rounded ; bearing a number of stout
hairs and short spines.

* This species is the black aphis found in spring on the spindle tree (Euonymus europaeus),
and later on beans, poppies, and many other plants. It has many synonyms, owing to its poly-
phagous habits, but the name given by Linnaeus in 1746 holds priority. The more important
synonyms are Aphis papaveris, Fabr.; Aphis euonymi, Fabr.; and Aphis fabae, Scop. For other
synonyms see Theobald, 1912.

+ The measurements given are total length of body excluding the cauda, and the greatest width
of the abdomen. The size is very variable, depending largely on food and temperature conditions.

¢ Accessory eyes (Berlese, “‘ Gli Insetti’’) ; ocular tubercles (Baker, 1920).

§ The “ unguis”’ (Baker, 1920), or “‘ processus terminalis’’ (Tullgren, 1909), of the terminal
segment is here considered as being part of the distal segment.

(2416) F

82 J. DAVIDSON.

First Larval Instar has 4-segmented antennae, with sensorium on segs. 4 and 5 ;
tubercle-like cornicles ; small obtuse cauda and stout legs.

Second Larval Instar has 5-segmented antennae.

Fig. 1. Aphis rumicis, Linn., fundatrix: (1) dorsal view ;

(2) antenna; (3) rostrum; (4) posterior end of venter, a.p.,

anal plate, g.p., genital plate, c, cauda; (5) tarsus of third
leg, , empodium or pad ; (6) cornicle.

2. APTEROUS VIVIPAROUS FEMALE (fig. 2). Average size 2-5 mm. by 1-6 mm.

Body elongate oval; colour variable, black to olive-green, often with irregular
darker pigmented areas over the abdomen ; small hairs scattered over the body.

Fig. 2. Aphis rumicis, Linn., apterous viviparous Q:
(7) dorsal view; (8) antenna ; (9) rostrum; (10) cauda,
dorsal view, a.p., anal plate, g.p., genital plate; (11)

cornicle ; (12) tarsus of third leg, p, empodium. .

BIOLOGICAL STUDIES OF APHIS RUMICIS, LINN. 83

Head: Eyes black, with prominent accessory eyes. Antennae six-jointed ; seg. 1,
apical portion of seg. 5 and proximal portion of seg. 6 black, remainder of a paler
colour ; seg. 6 (including processus terminalis) the longest, about equal to 4 and 5
together ; 3 about three-fourths the length of 6, longer than 4; 4 slightly longer
than 5; 1 and 2 subequal in length ; a single subapical sensorium on seg. 5; a
compound sensorium on seg. 6 ; a few hairs on all segments. Rostrum normal, with
a few hairs on the segments.

Thorax with a prominent prothoracic tubercle on each side. Legs black ; tibiae
and proximal portion of femora paler ; segments bearing stout hairs, especially the
tibiae.

Abdomen with two lateral tubercles on each side and sometimes one or two small
indefinite tubercles. Cornicles black, tubular, imbricated, slightly tapering distally,
varying in length, but usually about one and one-third to one and one-half times the
length of the cauda as viewed from dorsum. Cauda with distal half slightly spoon-
shaped, black, clothed with short stout bristles and several long curved hairs. Anal
and genital plates as in fundatrix.

3. WINGED VIVIPAROUS FEMALE (fig. 3). Average size, 2:4mm. by 1-3 mm.

Body: Head and thorax black to brownish black; the abdomen varying from
dirty brownish black to olive-green, usually with irregular darker pigmented areas on
the abdomen. Small hairs scattered over the body.

Fig. 3. Aphis rumicis, winged viviparous 9: (13) dorsal

view ; (14) antenna; (15) cauda, dorsal view, a.p., anal plate,

§-p., genital plate; (16) tarsus of third leg, p, empodium
or pad ; (17) cornicle.

Head black. Eyes black, with prominent accessory eyes. Antennae dirty brown
to black, varying in length, about two-third length of body; seg. 3 slightly longer
than 4; seg. 4 slightly longer than 5; 1 and 2 subequal; subapical sensorium on
seg. 5, and a compound sensorium on seg. 6; a varying number of about 12-18
subcircular sensoria distributed over seg. 3 and none or 1-4 over seg. 4; segments
3 to 6 imbricated. Rostrum dark towards distal end ; normal.

(2416) r2

84 J.. DAVIDSON.

Thorax with two prominent lateral tubercles on prothorax. Wings normal. Legs
somewhat longer than in a.v. 9, otherwise similar.

Abdomen varying in colour from dark velvet-black to olive-green, usually with five
irregular pigmented areas along the lateral dorsal area and irregular transverse areas
segmentally arranged; lateral tubercles prominent. Cornicles black, varying in
length, usually about one and one-half times length of caudaas viewed from dorsum,
imbricate, tubular, slightly tapering distally. Cauda not so large as in a.v. 9, other-
wise similar. Anal and genital plates as in a.v. 9.

4. SEXUPARAE.

(a). Male-producing sexuparae, apterous, resembling apterous viviparous female.

(5). Female-producing sexuparae, winged, resembling winged viviparous female,
but somewhat larger in size.

5. MALE (fig. 4). Average size, 1-4 mm. by 0:7 mm.

Body smaller than in w.v. 2, narrower and more tapering distally ; seen from
dorsum it appears to be shining black, but the abdomen is often very dark green ;
small hairs scattered over the body.

Fig. 4. Aphis rumicis, 3: (18) dorsal view; (19) antenna;

(20) rostrum ; (21) cornicle ; (22) tarsus of third leg ; (23) posterior

end of venter, showing genital armature, a.p., anal plate, c, cauda,
cl., claspers, p.s., penis sheath ; (24) cauda, dorsal view.

Head black. Eyes large, black; accessory eyes prominent. Antennae about
two-third to three-quarters length of body, black, but sometimes paler ; segments 1
and 2 subequal; 3 shorter than 6; 4 slightly shorter than 3; numerous subcircular

sensoria on segments 3, 4 and 5, anda compound sensorium on 6; a few hairs on all
segments. Rostrum normal.

Thorax black and shining ; prothoracic tubercles prominent. Legs slender, black,
with greater part of tibiae and femora paler ; hairs on all segments, especially tibiae.

Abdomen varying, almost black to dark green, black along lateral margins, irregular
patches of darker pigmented areas more or less segmentally arranged ; two lateral

BIOLOGICAL STUDIES OF APHIS RUMICIS, LINN. 85

tubercles present on each side. Cornicles dark to black, short, small, tubular, im-
bricated, a little longer than the cauda viewed from the dorsum. Cauda smaller than
in w.v. 2; covered with short spines and several long hairs; distal portion black.
Anal plate black. Genital plate black, bearing two dark claspers (gonapophyses)
clothed with spines and stout hairs. Penis sheath paler.

6. Oviparous FEMALE (fig. 5). Average size, 1-6 mm. by 0-9 mm.
Body small and narrow compared with the a.v. 2, tapering posteriorly ; dirty

brownish black to dark green in colour, often of a dark green velvety appearance ;

short hairs distributed over the body.

Head black to dark green. Eyes small, black; accessory eyes small. Antennae
about two-thirds length of body ; pale dirty grey, segments 1 and 2 and distal portion
of 4 and 5 darker; seg. 6 about equal in length to 3, 4 and 5 together; 3 longer
than 4 or 5; 4 slightly shorter than 5; a compound sensorium on seg. 6 and a sub-
apical sensorium on 5; a few hairs over each segment. Rostrum normal, dark on
distal portion, with fewer hairs on the segments.

Thorax with lateral tubercles on prothorax prominent. Legs relatively short and
stout ; of a dirty greyish colour, with coxae, trochanters and tarsi darker ; hairs on
all segments, especially on tibiae; tibiae of third pair of legs much swollen, and
possessing numerous irregular roundish light-coloured areas (? sensoria) over the
whole length.

Fig. 5. Aphis rumicis, oviparous @: (25) dorsal!
view ; (26) antenna; (27) rostrum; (28) tibia of third leg ;
(29) cauda, dorsal view; (30) posterior end of venter, a.p.,
anal plate, g.p., genital plate, c, cauda; (31) cornicle.

Abdomen dark green, with occasional lighter-coloured areas; two tubercles on
each side. Cornicles quite small, black, imbricated, tubular, a little longer than the
cauda as viewed from the dorsum. Cauda small but stout, black on distal portion.
Anal plate black. Genital plate bilobed, black, and covered with long hairs.

Ova, when first laid, somewhat greenish, but rapidly becoming black and shiny on
exposure to the air. Average size, 0-5 mm.

(24.S5—B)

86 J. DAVIDSON.

Il. Life-History of Aphis rumicis.

The following account of the life-history of this species is based upon extensive
breeding experiments and observations in the field. Experiments were carried on
during 1913 and the early part of 1914, and continued during 1920.

The Aphids were reared from eggs on Euonymus, and transferred to broad beans
as the summer host, the winged remigrantes or sexuparae of the later generations
being transferred back to Euonymus. The plants were grown in pots in a large open
glasshouse and kept covered with muslin bags, and observations on the different
generations of the Aphids were periodically recorded. It is hardly practicable, for
reasons of economy, to publish the observations made on the long series of plants
infected, but the records of certain plants will be given in later sections of these
biological studies. Itis from the data derived from these observations, together with
contemporary observations in the field, that the life-cycle has been worked out.

The ova hatch out in spring (March and April) on the winter host, Ewcnymus
europaeus,* giving rise to the larvae of the fundatrices. In 8-10 days after birth, in
favourable temperature conditions, the fundatrices become adult and begin to pro-
duce parthenogenetic viviparous young, which become adult about 10 days after
birth. This is the first viviparous generation, and it may consist of a mixed progeny
of both alate and apterous viviparous females, or of the latter only. The a.v. 9 9
produce on Luonymus the second viviparous g generation, which may consist of w.v. 2 9
and a.v. 9 9, the former being generally in the majority, The a.v. 2 2 produce a
further generation, consisting either entirely of w.v. 2 9, or of a mixed progeny of
w.v. 2 9 and a.v. 9 Q, the w.v. 2 9 being in the SEHD

There is a tendency for the a.v. 2 9 of these early generations on Euonymus to
produce w.v. 2 Qin the majority. These w.v. 9 9 are the winged migrants from the
winter host to the intermediate or summer hosts, such as beans, poppies, etc. It is
owing to this tendency that the a.v. 2 9 on Euonymus eventually die out, and as the
season advances the tree becomes free from the aphis. Some of the w.v. 9 Q may in
some cases remain a short time on Evonymus and produce young on it, but owing to
the innate desire of the winged forms to migrate (a marked feature of the winged forms
when the Aphids are reared in captivity), they soon leave the Euonymus and fly
to the intermediate hosts.

The generations of individuals from the fundatrices on Euvonymus are the fundatri-
geniae, “the a.v. 2 9 being the fundatrigeniae apterae and the w.v. 9 9 the fundatri-
geniae alatae. ince latter are the winged migrants (migrantes), which fly to the
intermediate hosts, being the mothers of the succeeding viviparous generations.
The generations of individuals from the migrantes on the intermediate hosts are
called the alienicolae, the a.v. 2 9 being the alienicolae apterae, and the w.v. @ 9
the alienicolae alatae.

The intermediate hosts are numerous, but of the cultivated plants beans are specially
favourable, and afford the best stimulus to rapid reproduction. The migrantes give
rise to the first viviparous generation on the intermediate host, the individuals of
which area.v. 9 9. These produce a further generation, which may consist ofa.v. 2 9
or a mixed progeny of w.v. 2 Qanda.v. 9 9. The former may remain a little time on
the same plant and produce young, or may fly to other plants, either of the same kind
or other hosts, and produce a further generation. My experiments show that the
tendency is for w.v. 2 9 to produce a.v. 9 9, and for a.v. 2 9 to produce either
a.v. 9 9 or a mixed progeny with a varying percentage of w.v. 9 9. This would

* The spindle-tree is undoubtedly a winter host of Aphis rwmicis, but considering the local
distribution of Euonymus in Britain, it is highly probable that there are other winter hosts.
Gaumont (1913) found all stages on E. japonicus. Mordwilko (1907) found Aphis euonymi in
spring on Viburnum opulus at Bjelovesh. It was also found on Viburnum opulus by Kaltenbach
(Aphis euonymi) and by Passerini (Aphis papaveris). It is evident that further research is
greatly needed in order to ascertain the common winter hosts of this abundant Aphid.

BIOLOGICAL STUDIES OF APHIS RUMICIS, LINN. 87

appear to be an adaptation of the species to its polyphagous habits. Thus when the
w.v. 2 Q alight on a new plant a good colony is established on that plant by a.v. 2 &
being produced. On the other hand, the lability to overcrowding as the plant
becomes heavily infested is overcome, and the wider distribution of the species to
other host-plants is ensured by the production of winged forms in due course.

Towards the end of summer, after a number of agamic generations have been passed
through on the intermediate hosts and at a time when suitable intermediate hosts are
becoming scarce, there are produced winged viviparous females, which are physio-
logically specialised, but morphologically resemble the other alienicolae alatae.
These are the sexuparae alatae or remigrantes. They fly back to the winter host, on
which they produce true oviparous @ 9.

At about the same period winged g 3 are produced from certain of
the alienicolae apterae (sexuparae apterae) on the intermediate hosts and fly tothe
winter host, where the sexual 9 9 are fertilised. Fertilised eggs are then laid on
the winter host, near the buds, or in crevices in the bark of the older branches.
These over-winter and hatch out in spring, producing the fundatrices.

The alienicolae apterae on the intermediate hosts gradually die out, owing partly
to the tendency to produce w.v. 22 or winged sexuparae and gg, and
partly to unfavourable conditions.* By confining the Aphids to broad bean plants,
sexual forms were produced on these plants in due course.

In my experiments it was found that the alienicolae alatae of any generation could
be transferred back to Euonymus, on which plant they produced young, and eventually
in succeeding generations both sexual 3 ¢ and sexual 2 2 appeared. Further, it
was found that even if the Aphids are bred in successive generations on Euonymus,
sexual $ g and © 9 will appear in due course. It should be noted, however, that
young growth was ensured on the Exonymus bushes by cutting them back. Males
were first noted on 10th August 1920.

Similarly on beans, on which intermediate host the Aphids were bred continuously
from May, several ¢ 3 appeared in due course, and in some cases oviparous
2 © were also found. The 3 3 are produced by alienicolae apterae toward the
end of summer on the intermediate host-plants, and the oviparous 2 Q by
the physiologically specialised alienicolae alatae (sexuparae alatae) on the winter
host, at about the same time. The male-producing sexuparae are thus a.v. @ 9.
One and the same mother may produce sexual males and sexuparae alatae, but
the sexuparae alatae only produce sexual females.

It seems evident that in the adventures of migration the alienicolae alatae of any
generation may alight on the winter host (Ewonymus) and produce young, resulting
eventually in colonies consisting of all stages, namely, a.v. 2 9, w.v. 29 9, § gd and
2 9. Mordwilko (1907) found all stages on Ewonymus in Warsaw at end of September
1894. He also found a colony of a.v. 2 2 on Viburnum opulus in Warsaw at the end
of September.

Similarly, sexuparae alatae alighting on intermediate host-plants may give rise to
sexual 2 9, which might thus be found together with colonies of agamic forms and
sexual males. This would explain the finding by Theobald (1912) of sexual 2 &
ovipositing on Rumex, by Gaumont (1913) on sugar-beet in October 1913, and by
Malaquin and Moitié (1914) on haricot beans in October 1913.

* It seems probable that during a mild winter agamic forms may persist throughout on certain
plants and carry on agamic reproduction normally in the following year. Davidson (1914) found
a colony of apterous agamic females on 30th January 1913 on Euonymus at Richmond. The
tree was taken into a greenhouse, and agamic reproduction was carried on normally throughout
1913. Several cases of long-continued parthenogenetic reproduction have been observed. The
question will be discussed in a later section dealing with the appearance of sexual forms. It may
be stated here that I have carried on a parthenogenetic strain throughout winter in a warm green-
house, winged sexuparae (which produce sexual 99), sexual Gg and parthenogenetic a.v. oye)
being produced in each generation from September to May.

t Mordwilko (1907) was not able to obtain females on intermediate hosts.

88 J. DAVIDSON.

During the autumn, when the intermediate hosts are mostly unfavourable for the
Aphids, the winter hosts, such as Euonymus, offer the most favourable food con-
ditions and the greatest chances of survival. From the nature of Rwmex and other
intermediate hosts under winter conditions and the difficulty in ensuring food for the
young larvae immediately they hatch out from the eggs in spring, these cases, I think,
must only be considered as casual winter hosts. The sexual forms may develop in
several generations. In fact, in one series of experiments under favourable conditions
of food and temperature, sexual forms appeared in five succeeding generations.
Weather conditions and the dying down of the intermediate hosts are very important
factors in limiting the length of period over which sexual forms are produced. These
questions will be discussed more fully in a later section dealing with the appearance
of the sexual forms.

The life-history of Aphis rumicis may be illustrated by the following diagram :—

©= Ova. WE = Fundatrices. @=Apterous viviparous females. O = Winged viviparous females.

@ = Migrantes. O= Alate sexuparx2 (Remgrantes). of = Sexual males. e) = Oviparous females.

Fig. 6. Diagram illustrating the life-cyle of Aphis rumicis. The line

AB divides the circle into two halves, the winter host being represented

on the left and the summer hosts on the right half; the sectors indicate

the months, and the dotted concentric lines a varying number of generations
occurring in July and August.

BIOLOGICAL STUDIES OF APHIS RUMICIS, LINN. 89

References.
Davipson, J. (1914). Ann. Appl. Biol. i, pp. 118-141.
Gaumont, M. L. (1913). C.R. Acad. Sci. Paris, clvii, pp. 1092-1094.
LINNAEUS, CAROLUS (1746). Fauna Suecica, p. 258, No. 979.
MorDWILko, A. (1907). Biol. Centralb. xxvii, p. 529 et seq.
THEOBALD, F. V. (1912). Journ. Bd. Agric. xix, pp. 466-476.

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THE NATURE AND FUNCTION OF THE CAUDAL TUFTS OF MALAYAN
ANOPHELINE LARVAE.

By W. A. LamBorn,
Malaria Bureau, Federated M alay States.

It has been found difficult to understand how such a fragile creature as an
Anopheline larva could withstand the force of a strong current of water. In the
following article it is proposed to explain the phenomenon on anatomical grounds.

The writer’s previous experiences in Central Africa, where the seasons are far more
definite than in the Federated Malay States, had shown that in the dry period, when
there is often an entire absence of other possible breeding-places, the larvae of
Anophelines are to be found in among the vegetation fringing the banks of the great
rivers ; hence, for instance, the notoriety as regards malaria of the whole extent of
the Shire valley, in N yasaland.

It was decided in June 1920 to make a tentative examination for larvae of the
Klang River and its affluents in the vicinity of Kuala Lumpur, F.M.S., though there
was not at that time any great expectation of being successful in the search. It is
true that Dr. Hacker had in 1918 demonstrated the presence there of the larvae of
A. maculatus, Theo., and A. aconitus, Dén. ; but apart from the numerous breeding-
places afforded elsewhere, swamps, grassy pools, muddy pools, drains, and excavations
of various sorts, the heavy rains at frequent intervals during the previous three months
had ensured the flow of a volume of water so enormous and a current so swift as to
make it appear impossible that an object so fragile as a mosquito larva could maintain
itself therein.

A few Anopheline larvae—A. subpictus, Grassi, var. vagus, Don. (A. rossi, Giles,
var. indefimitus, Ludl.), A. barbirostris, Wulp., A. hyrcanus, Pall. (A. sinensis, Wied.),
A. acomitus, Dén., A. maculatus, Theo., A. leucosphyvrus, Dén.—were discovered, and
even with the river running strongly, could constantly be found in among the stems of
the reeds and long grass at its sides. At no time were they numerous, and careful and
diligent search was necessary to obtain them in a rapid current. The significance of
their presence would appear to lie, not in their numbers, but rather in their ability
to maintain themselves in such a situation, from which in due course the adults can
diffuse themselves over the surrounding country, selecting at will, possibly at another
season, breeding-places more auspicious to the welfare of their larvae.

The question which then arose was: How do the larvae manage to maintain them-
selves without being swept away? For not only in the river, but in other running
water are the larvae of certain species constantly to be found.

A. maculatus larvae, for instance, can invariably be found in a grassy drain beside
a railway embankment just outside the town, in which a considerable volume of water
is usually flowing. A few A. karwari larvae were once found by the writer at Sungei
Besi, eight miles south of Kuala Lumpur, apparently quite at ease at the side of a
pot-hole through which the water was literally rushing from a higher level. The
larvae of A. aitkeni, James, almost invariably favour jungle streams.

Some observations made in this connection so long ago as 1911 by Dr. Malcolm
Watson, and recorded in his book “‘ The Prevention of Malaria in the Federated M alay
States ” (p. 105), are of no small interest. Dr. Watson expresses the opinion that the
ova of A. maculatus (wilmort) are laid in the shallowest possible water, usually at the
head of a ravine, and the paragraph discussing further the distribution of the larvae
is as follows :—

“From these springs the larvae are carried down by the streams, but they cannot
be entirely washed out even by the strongest currents or rains. I have found them in

92 W. A. LAMBORN.

a drain after a two-inch shower. I have watched them at play in a clear pool at the
foot of a rock down which water was flowing with considerable force, since the rock
sloped to the pool at an angle of forty-five and its face was a foot long. The current
of water was still further increased in strength by the rock being funnel-shaped, and
all the water coming down the face was gathered into a solid stream as it entered the
pool. The larvae were playing not exactly like trout, head to stream, but were floating
round in the current, and every now and then one would swim right into the stream,
up it for a short distance, and then hang on the side of the apparently bare rock in the
full strength of the current.”’ Further on (p. 116) he remarks that “‘ the larvae have
the power of attaching themselves to objects.”’

Anopheline larvae, at all events such as live in moving water, do not, as a rule, go
in search of food. They rest at one place, brushing constantly the minute particles
carried past into their mouths, consuming such as they wish, and ejecting the rest. It
is hardly to be expected that a larva undergoing extremely rapid growth and develop-
ment, on what would appear to be a precarious method of obtaining food material,
could afford to waste any of its energy through having to keepits position by strong
and constant muscular effort. Some sort of mechanism involving a minimum
expenditure of energy, and not a means similar to the muscular effort that enables
a fish to remain at one spot in a stream, was to be looked for. This is found in the
dorsal pair of tail fans, the true function of which is now pointed out for the first
time, so far as the writer has been able to ascertain.

Patton and Cragg, in their “‘ Text Book of Medical Entomology ”’ (1913, p. 200),
describing the ninth segment of an Anopheline larva, state that “ the dorsal border is
furnished with two pairs of long feathered hairs, which are directed backwards as a
tail. The ventral surface bears wo rows of feathered hairs arising in the middle line
from an elongated and raised area of thick chitin. The two rows are set very close
together, so that when examined in side view they appear as one, and hang down at a
right angle to the long axis of the body as a sort of fin. Each hair is articulated into a
little round pit in the chitinised area.”’

Giles in his work “‘ Gnats or Mosquitos ”’ (second edition, p. 46), describing the larva
of a Culex, states that on the last segment, “‘ on either side, but originating in frent of
the anal tubercles, are a pair of large dense tufts of compound hairs, which are
employed in swimming in the same way as a fish’s tail, and are so arranged as to form
an expansion of similar shape.’’ In his description of an Anopheline larva (p. 60)
he states that “‘ the last segment carries four anal tubercles, which, as well as the tail
fans, are rather less developed than in Culex.”

The American authors—Howard, Dyar and Knab—in their “‘ Mosquitoes of North
and Central America and the West Indies,”’ do not give any special description of
these tufts in Anopheline larvae.

From the foregoing it is evident that no structural differentiation of the two sets
of brushes in larvae had been noted. In the larvae of Malayan Anophelines the
ventral tufts are as described. But with an objective even of such low power
as 2 in. marked differences are to be seen in the dorsal group. For the
purposes of description the larva of A. maculatus at its last moult may be taken as
presenting appearances more or less typical of the majority (fig. 1). In this the dorsal
aspect of the ninth segment is furnished at its extremity with two brush-like structures
on either side, each just to the outer side of the mid line. The upper and more internal
brush consists of a short plume of feathered hairs, which are straight, scanty, and on
one side almost double the length of those on the other. Most of the hairs taper
gradually and terminate in a sharp point, but one or two of the longer ones may show
a terminal hooklet so smallastorequirea } in. objective for its detection. The second
structure, just to the lower and outer side of the former, consists of a leash of six
stout bristles, five of which are unbranched. They are considerably longer than any
in the other brush, there being a slight progressive increase in the length of each, so

;

THE CAUDAL TUFTS OF MALAYAN ANOPHELINE LARVAE. 93

that the longest is about a quarter as much again the length cf the shortest. All are
of about the same calibre, which is practically uniform from end to end, and each of
the five is recurved at itsextremity, so as to form a rounded hook. The sixth bristle
differs from the other five in being considerably shorter and in bearing seven lateral
filaments, all of similar calibre but of different lengths, the two longest showing hooks.
Not one of the bristles is perfectly straight, each showing a curve first upwards and
then gradually in the reverse direction, more sharply towards the extremity. The
ventral brushes consist of separate tufts of feathered filaments, arranged fanwise,

Fig. 1. Terminal segments of larva of 4 nopheles maculatus, Theo.

tapering gradually towards the extremity, which is pointed. They are apparently of
less brittle material than the dorsal bristles ; for whereas these often have the hooks
broken off, or may be broken short when the larva has been much shaken about, the
central filaments seem not to suffer such damage.

The hooked bristles are present throughout larval life. In the newly hatched larva
there are two only on each side, and the number increases at each moult. The same
structures are to be seen without any modification of form in the larvae of the following
Anophelines :—A. karwart, A. hyrcanus (sinensis), A. kocht, A. ludlowt, Theo., and
A. subpictus (rosst) and its var. A. vagus (indefinitus). In A. fuliginosus, Giles, A.
barbirostris, A. umbrosus, Theo., A. tessellatus, Theo., and A. aconitus, there are, as a

Fig. 2. Terminal segments of larva of Anopheles asiaticus, Theo.

rule, five main hooked bristles only; in A. albotaeniatus, Theo., var. monianus,
Stant. & Hack., there are, asa rule, five, and the upper of the dorsal tufts shows
considerable reduction in size. In the case of A. aitkeni, James, and A. asvaticus,
Theo., there are marked differences ; in aitkeni there are six main hooked bristles, one
of which shows a branch almost as long as itself and also hooked, so making an
additional main limb ; moreover, the upper of the two dorsal tufts shows many more
fine hooks at the extremities of the constituent hairs than in other species, doubtless
in adaptation to its mode of life, the species breeding in mountain streams, Mm
asiaticus (fig. 2) there are only two, or sometimes three, hooked bristles, which are

94 Ww. A. LAMBORN.

attenuated as in Culex and often crooked, and show hooks so diminutive as to make
the determination of their presence difficult. In this also the upper of the dorsal tufts
is reduced to a mere bristle clothed with a few sparse hairs.

It was of interest to learn from Col. Alcock, of the London School of Tropical
Medicine, that these structures are present also in two of the British species, and that
in the third, breeding in situations comparable with those favoured by A. asiaticus
in the Federated Malay States, they are, as in that species, absent. The paragraph in
his letter, dated 10th August 1920, which he has kindly allowed me to quote, is as
follows :—

“On looking at the larvae of the three British species (after reading your letter),
I find it to be as you describe in A. maculipennis and in A. bifurcatus, but not in A.
plumbeus, that breeds in holes of trees. Now I understand how the larvae of the former
two species anchor themselves at right angles to the sides of the aquarium.”

Were these structures used for swimming, one would expect to find them at their
maximum development in larvae which move more actively than most Anophelines
in search of food, but this is not the case. Stegomyia probably owe their success as
a race of mosquitos, not merely to their being able to breed in almost any situation
invariably in stagnant water in which insect enemies are commonly absent, but to the
activity they are able to develop in their search, both at the surface and at a depth,
for the scanty and miscellaneous food material such places often afford. Placed in
a bowl, they may be seen moving constantly, head down, over any surface on which
food material may be obtained ; they never remain passive at the side, content to
take such food as comes along, as some Anopheline larvae do. In these larvae the
various caudal structures are least developed, being represented by a much simpler
arrangement of hairs. There are in the last ecdysis three dorsal and three ventral
hairs only on each side, all relatively longer than in Anophelines (except in A. aszaticus),
perfectly straight and tapering gradually so as to end in a fine needle-point.

Anopheles larvae may be called on at times to make a sudden and very rapid
movement ; this is essential to enable them to escape their enemies, and it is effected
by all these larvae, which swim backwards, at all events when on the surface of the
water, by sudden contractions of the body, as in the case of Stegomyia. But it is
essential for such as live in moving water that the movements should be to some extent
objective, towards some support, so that they may avoid being swept away. It is
suggested that direction is attained by means of rudder-like action of the ventral
fans, also much more highly developed than in Stegomyia, which, furthermore, enable
them on occasion to turn sharply so as to attach themselves to any support by means
of the hooks of the dorsal brushes, which have just been described.

The function of the hooks was very readily determined by affording larvae in a glass

dish, a small twig as a rough object of support. The larvae of all species except
A. asiaticus, especially those commonly found in moving water—A. maculatus and
A. karwan for instance—invariably take advantage at once of such a support,
resting at a right angle to it. Even with the unaided eye, when the larva is at rest
against the support, it can be seen that the caudal tufts are so arranged that the ventral
group and the inner dorsal brush remain in direct line with the body, the latter just
touching the support, and that the dorsal leash of hooked filaments is directed out on
either side at an angle of about 45 degrees to the body. If the supporting object is
slightly submerged the brushes are directed down, obliquely or vertically, but still
occupy the same relative position. By the aid of a microscope it can be determined
that the hooklets are twined round any rough point, so neatly sometimes as almost to
tempt one to suspect a real tendril-like action.

If the supporting body is dragged through the water, larvae may be drawn along
still clinging to it, and the support may be rather violently moved to and fro, the

THE CAUDAL TUFTS OF MALAYAN ANOPHELINE LARVAE. 95

attached larvae then swaying backwards and forwards from the point of attachment.
Larvae are able readily to attach themselves to moving objects.

In this way, then, the larva is able to avoid being swept away, and it can also employ
both sets of hooks together at one point, thenresting more or less parallel to the support-
ing object, its head down current, a measure of value doubtless when it has to sustain
itself against a greater pressure of water than it could withstand in any other position.
When the larva is floating unsupported against any object, the caudal fans are approxi-
mated and directed back, and in this position it was noted occasionally, especially in
A. maculatus, A. indefimitus and A. aitkent, that the hooks from the bristles of opposite
sides may be interlocked.

Were any confirmation as to the function of the dorsal tufts needed, it is afforded
by the study of these structures in A. aithent and A. astaticus. In A. aitkent, a regular
breeder in mountain streams, the number of hooks available for the support of the
larva is at least double, since both dorsal tufts bear them. In A. astaticus, a breeder
in bamboos, the structures show a very definite tendency to atrophy, a fact suggesting
that the very special sort of habitat selected for the larvae is by no means a very recent
choice on the part of the female parent, an inference supported by the size of the
prunitive larval eye, which is very much smaller than in the larvae of any of the other
species of the group, and by the absence of the group of pigment spots arranged in
crescentic form, which in other species represent the developing compound eye. It
is interesting to note in the case of the bush-breeding Anophelines and in Stegomyia
(also a breeder by preference in dark places), an approximation toa similar condition.*
The presence of any hooks at all in the case of A. subpictus var. vagus, and others which
breed by choice in the still water of muddy pools, is doubtless to be explained by recent
modification of breeding habits ; for until the advent of the white man to this country
and the subsequent great economic development, there must have been comparatively
few such breeding-places available.

Though these hooks must be the chief means of support to larvae, it is certain that
the young larvae of A. maculatus, A. karwari and A. aconitus are able to attach them-
selves to an object for a short period of time by their mouth-parts. It was necessary
in the course of some breeding experiments in connection with these species to transfer
young larvae from one porcelain bowl to another, and it was repeatedly found that
some with their heads applied to the surface of the bowl could resist withdrawal into
a glass tube, and that if a current of water was then directed on them, they were not
readily dislodged, swaying to and fro at the point of attachment. The porcelain being
glazed, it would appear as if they had some power of attachment thereto by suction,
no hooklets being present on the mouth-brushes.

The supporting hooks found in the case of the larva would benefit the several
species but little, if there were not some sort of counterpart in the case of the pupa.
The usual text-book description of the way in which a pupa evades danger is that it
does so by diving and keeping itself submerged. Its specific gravity being less than
that of water, it remains below either by getting under some object, or by holding fast
by clasping it between its thorax and its flexed abdomen. This appears to be
true of certain of the Anophelines, but not of all. It was recognised, for instance, by
the writer early in the course of a study of A. vagus in its muddy pools that the pupa
is able to sustain itself with its tail, apparently just simply applied to any portion of

* In this connection it is noteworthy that, whereas in the fully-grown larvae of A. hyrcanus
and A. barbirostris (open-country breeders) the pigmented elements of both larval and imaginal
eyes reach a maximum development, in the case of A. umbrosus, the remaining Malayan repre-
sentative of the Myzorhynchus group of Anophelines (a breeder by preference in jungle), they are
much reduced in size, especially in the crescentic eye, in which indeed they are so pale that the
determination of their presence is difficult.

96 W. A. LAMBORN.

mud on the bottom. Occasionally a pupa may be seen rising slowly to the: surface
with material from the bottom dangling from the caudal fins. It has been able so to
attach itself by means of a pair of hooklets, one springing from the free margin of each
fin, and actually an extension of the thick chitinous bar, which forms a backbone
supporting the fin.

If one studies vagus pupae in a test-tube, one finds that for the purpose of maintain-
ing themselves below the surface, one of three common methods may be adopted,
according to the nature of the supporting material available. First, by preference
a pupa will get beneath the supporting object ; secondly, if the object is of convenient
size it may clasp it with its anteflexed abdomen ; or thirdly, it may attach itself by
the caudal hooks, then rising above the support. Occasionally a pupa of vagus
has been seen upside down, attached to a rough support by means simply of one
respiratory trumpet, showing how slight must be the difference in specific gravity
between itself and the water in which it is found. Though the hooks (fig. 3) are present

Fig. 3. Terminal segments of pupae of: (a) Anopheles

subpictus, Grassi; (b) A. subpictus var. vagus, Don.; (c)

A. asiaticus, Theo. ; (d) A. hyrcanus, Pall. ; (e) A. tessellatus,
Theo. ; (f) A. karwari, James.

in the pupae of subpictus (vosst), maculatus, karwari, aconitus and fuliginosus, they are
represented in sinensis, barbirostris, umbrosus, kochi, tessellatus, asiaticus, and, curiously
enough, in aitkent also, by a mere filament, and in these latter species, the pupa attaches
itself to an object by one of the two ways described first, or, in the absence of any
support, remains below by swimming efforts, renewed directly it tends to rise.

The form of the hook varies a little in each species. In all it consists of a stout
bristle, which after a short straight course becomes recurved in the direction of the
dorsal aspect of the pupa, terminating in a rounded hook. The bristle itself is of brittle
material, for often one or both may be broken off short in pupae which have been
roughly dealt with.

THE CAUDAL TUFTS OF MALAYAN ANOPHELINE LARVAE. 97

Conclusions.

1. The larvae of certain Anophelines, particularly A. maculatus, A. karwart and
A. aitkent, occur in streams so swift that their presence would have appeared
impossible.

2. Some mechanism must be looked for which would enable larvae to maintain
their position in the water.

3. A pair of the dorsal tail brushes, terminated by hooklets, provides this mechanism;
and there is a counterpart, in the case of certain pupae, in the form of a pair of hooks
terminating the paddles.

The writer’s acknowledgments are due to Mr. M. C. Chuen, Laboratory Assistant
at the Malaria Bureau, Kuala Lumpur, for the care and skill he has exercised in
preparing from living specimens, the drawings which illustrate this note.

Kuala Lumpur, F.M.S.
Sept. 1920.

(2416) G

FLAX CATERPILLARS IN KENYA COLONY, WITH SPECIAL
REFERENCE TO THE ‘LIMITATIONS. OF THE. ROPING
METHOD OF COMBATING THEM.

By F. W. Dry, M’Se;,
Recently Assistant Government Entomologist, Kenya Colony.

Caterpillars on flax in Kenya Colony are farm pests of the first rank. They belong
to more than one species, but the one most commonly reared from material from the
field is Phytometra (Plusia) orichalcea, ¥.; Heliothis obsoleta, F., is also met with.

Caterpillars have been reported from all the chief flax-growing districts of the
country: the Kikuyu district, including Kabete, Kyambu, Limuru and Thika, and
the districts of Nakuru, Lumbwa and the Uasin Gishu. Near Kericho, which is
wetter and colder than Lumbwa, from farms up to within a distance of six miles
from those of the Lumbwa area, which have suffered badly, caterpillars have been
reported, but no serious damage has been done. Mr. T. J. Anderson, the Government
Entomologist, has taken the moths of Phytometra orichalcea in the Trans-Nzoia
district far from European cultivation.

The caterpillars have a wide range of food-plants. In East Africa they have been
found to attack, in addition to flax, Canadian Wonder beans, potatoes, rape and
other crucifers, as well as weeds.

The attack on the flax crop may take place at any stage, from the time the crop
is only just above the ground up to the time of pulling. In its early stages the crop
may be eaten upcompletely. In the later stages the seed-bolls are especially attacked.
In a bad attack on the partly grown crop all the leaves and the tops of the stems
are consumed, all that remains being the stripped stalks. When the caterpillars are
less numerous a frequent form of damage is that the growing-point at the apex of

the plant is killed and several weak secondary stems are thrown out. This results
in shortening of the fibre and irregularity in the ripening of the seed. :

The damage caused by flax caterpillars has been very serious, and many fields have
been destroyed. There have been heavy losses in each of the three years 1918,
1919 and 1920, the worst reports having been received during the last year. Most
of the damage is done from May to August, following the long rains, in the main
growing season, but outbreaks have also occurred in the season following the short
rains, which are due from October to December.

The life-history of Phytometra orichalcea was worked out in the laboratory at Kabete
during the months of June to August, 1918, comparatively cool months. Eggs were
deposited on the leaves or stems of the plants provided. The cocoons were spun on
the host-plants, several leaves or stems being bound together. The caterpillars were
fed on the weed Galinosoga parviflora. Little variation was found in the length of
the life-cycle of different individuals. The average times for the different stages
were :—

Egg a, ae se ae Sar te: ey | SLOT days:
Caterpillar (hatching to spinning cocoon) <a POs ... 41 days.
Cocoon (spinning to emergence of moth) ais ee ... 30 days.

Eggs were laid a day or two after the emergence of the moths, making the length of
the life-cycle, from egg to egg, about twelve weeks.

The caterpillar stage lasting, as it does, about six weeks, there is some little time
for the detection of the pest while the caterpillars are still small. The large caterpillars
are quite conspicuous, but the younger ones harmonise so well with their surroundings
that unless a careful search be made they may easily be overlooked.

100 F. W. DRY.

At least eight species of parasites, some Hymenopterous, some parasitic flies, have
been reared from material from the field at Kabete.

Little success in controlling the caterpillars has been obtained by spraying. In
May 1919 a number of spraying experiments were carried out in which strong Paris
green did produce some results. The Paris green was used at the high rate of 3 lb.
to 40 gallons. This was combined with (a) soap, (b) resin-washing-soda sticker,
(c) lime, (d) lime and sticker. With these four plots were sprayed side by side, and
subsequently some small dead caterpillars were found, the caterpillars in the field
being mostly only about half-grown. Dead caterpillars could not be found on un-
sprayed control areas. About ten days after spraying there was a great difference
between all these four plots and adjacent control areas. On the Paris green plots
little damage could be seen as one looked across the field ; on the controls the cater-
pillars had retarded the growth considerably. A sharp line was seen between sprayed
area and control. Later, the weather being damp, the control areas caught up with
the sprayed parts of the field.

At other times spraying experiments with Paris green against larger caterpillars
have not met with success. One planter, who did not supply full data, reported
success with Paris green and lime, but the position is that one cannot recommend
spraying to planters.

The method of combating the caterpillars by “ roping’’ was devised by Mr. J.
McDonald, a Lumbwa settler, whose description is now quoted :—

“A rope, 40 yards long and about one inch in diameter, held by a boy at each
end, is dragged through the flax, the infested area being gone over from three to six
times a day. If necessary more than one rope is kept in action at once. The rope
should not be held too tightly. The boys use the same tracks each time and very
little mechanical injury will have been done to the crop. The method is continued
for several days, until the caterpillars have disappeared.”’

Here it is interesting to note, though the principle is not the same, that Dutt*
in India, speaking of the caterpillars of Phytometra orichalcea in peas, says, ‘‘ Dragging
a rope, moistened with kerosine and turpentine, over the crop drives away the
caterpillars. Even if they go into the neighbouring fields the crop is saved.”’

This roping method of Mr. McDonald’s has been tried by quite a number of flax-
growers. Their reports have varied. Some have said it was entirely successful,
others that it was no use. Success, however, was reported in a sufficient number of
cases to suggest that the method was of some value, and a week on farms in the
Nakuru district, in June 1920, gave the opportunity of applying the tests to this
method now described.

The method adopted was to obtain an index of the number of caterpillars in a
flax field where roping was in progress and then, four days later, to obtain another
index of the numbers present as a test of the result of roping. These numerical
estimates were made by walking slowly through the flax and counting the number
of caterpillars seen in five minutes, several such counts being made on each occasion.
The counts were made by the same people each time, in the same part of the field,
and at the same time of the day.

The results may be summarised as follows :—

Field No. 1.—The seed-bolls were just forming on the flax. Caterpillars were quite
numerous, Just a very few had formed cocoons. After four days roping with heavy
ropes about an inch in diameter the caterpillars were only a quarter as numerous as
when the first count was made. There were a few more cocoons than previously,

* Report of the Proceedings of the Second Entomological Meeting, held at Pusa on the 5th and
12th February 1917.

FLAX CATERPILLARS IN KENYA COLONY. 101

but the increase in their numbers was entirely insufficient to account for the reduction
in the numbers of caterpillars. Some may possibly have pupated in the ground (but
compare with Field No. 2).

Field No. 2.—This field was on the same farm as No. 1 and quite near to it. The
flax was nearly ready for pulling. The caterpillars, which were especially attacking
the seed-bolls, were rather less numerous than in Field No. 1, but they were rather
older and many cocoons had already been spun. The field was treated in the same
way as the other, but after four days no reduction was found in the number of
caterpillars.

Field No. 3.—In this field on a neighbouring farm the flax was in flower. The
caterpillars were about as numerous as in Field No.1. The field had been gone over
about six times a day, but instead of rope, reim (twisted hide) had been used, the
reim being much less heavy than the rope used on the first two fields. The reduction
in the number of caterpillars was about 35 per cent., as against about 75 per cent.
on Field No. 1.

Field No. 4.—As a check on the counts in the above three fields, counts were made
on another field, which had not quite reached the flowering stage, where the caterpillars
were not very plentiful. They were of fair size, but no cocoons were found. This
field was not roped. The number of caterpillars found after the four days’ interval
was almost identical with that found previously.

Field No. 5.—On this field of flax in flower there was a very bad outbreak. Many
of the caterpillars were in the last stage and cocoons were very plentiful. On part of
this field the flax had been badly stripped of leaves before the outbreak was observed,
and roping started, but caterpillars, while less numerous, were still present in large
numbers in other parts of the field, and in these parts not many cocoons were seen.
After four days of roping, with ropes about an inch in diameter, dragged through the
flax half a dozen times a day, the caterpillars were appreciably fewer than previously
on part of the field where the crop had not been seriously damaged, but the roping
had not prevented the badly stripped area from increasing in size.

In the last field, some observations were made in order to try to determine just in
what way the roping may bring about a reduction in the numbers of the caterpillars.
Dead or injured caterpillars on the ground are not objects which readily catch the
eye, but when a search was made some injured caterpillars were found. One large
caterpillar which could just wriggle a little was found, and an hour later it was unable
to move, being apparently dead. Another caterpillar, not very large, was found on
the ground, moving only just a little ; it did not make any attempt to climb up a
flax plant ; several small ants, one after the other, were seen to tackle it single-handed,
when it would wriggle, and each time they gave up the attempt ; but after an hour
and a quarter it was attacked by these ants in force and carried off. Other caterpillars,
some larger, some smaller, were seen to be attacked by ants successfully. In an area
of two square feet where particular search was made, six victims of ants were found.
One, which the ants had been watched carrying-off, was found an inch anda half below
the surface of the soil. Another, injured at the hind end, succeeded at the third
attempt in escaping from its assailants up a flax stem, which it climbed about three
inches, but there it hung helpless from a leaf. It was found that a big healthy cater-
pillar was able to escape, even when placed in an ant-run, but half-grown healthy
caterpillars were overwhelmed and carried off by a crowd of ants. Very small
caterpillars were several times found being carried by only a single ant.

Obviously, from the fact that great numbers of caterpillars are seen climbing up
the plants after being knocked off by the rope, the great majority escape after any
one passage of the rope, but if only a small percentage be mortally injured, or placed
at the mercy of ants, the cumulative effect would explain how the roping method
acts. Six per cent. of casualties each time the rope passes, is sufficient to explain tle
figures recorded for Field No. 1.

102 F. W. DRY.

Another explanation of the action of the roping method which has been put forward,
is that the caterpillars, becoming weary of being repeatedly knocked off the plants,
migrate from the field. A watch was kept for such migration from the field under
discussion, but caterpillars were only found crawling out of the field where the flax
had been stripped of leaves right up to the edge of the field. Where the flax offered
plenty of food, the caterpillars remained there. It seems probable that the roping
method, in so far as it does reduce the number of caterpillars, acts not by driving
them out of the field, but by bringing about their death in the field.

The conditions involved in the success or failure of this method seem, therefore,
to be :-—
(1) Kind of rope and frequency of roping.
A heavy rope has more effect than a light one. The more times the field
can be gone over ina day the better. If caterpillars are present in sufficient
numbers to do appreciable damage, once or twice a day would do little
good.

(2) Size of the caterpillars.
On small caterpillars the roping method has an effect both directly, by
injury, and indirectly, by placing them at the mercy of ants. On large
caterpillars there is little effect by either means.

(3) Numbers of the caterpillars.
Against an outbreak in which the caterpillars were very numerous, I doubt
the efficacy of this method, unless, possibly, roping were done more times
a day than would be feasible on a farm. Very little success has been
reported against bad outbreaks, and in the light of the field
observations just recorded this is not surprising. For,

(a) It appears that roping produces its effect by bringing about the death of
only a small percentage of caterpillars each time the rope passes.

(b) Unless the numbers of ants or other predators attacking the caterpillars on
the ground are increased by immigration, the larger the numbers of
caterpillars, the less potent proportionately will the ants be to reduce the
numbers of the caterpillars.

These facts, I believe, explain the different reports on the roping method—some of
success, some of failure—which have come to hand. If the caterpillars are big before
roping is started, if their numbers are very large, if the rope is too light, or if it is not
passed through the field often enough, failure may be expected. But if the cater-
pillars are small and not present in excessive numbers, roping, properly carried out,
will, I believe, meet the case.

It will thus be seen that there are distinct limitations to the roping method, and
it is, moreover, a laborious one. Undoubtedly, it will be replaced by something more
effective, and at present flax-growers are paying attention to various mechanical
devices invented by several settlers for removing the caterpillars from the crop, while
the search for other methods is being continued by the Division of Entomology.

103

THE RED SCALE, CHRYSOMPHALUS AURANTITI, MASK.,
IN KENYA COLONY.

ay EW. Dry, Msc.,
Recently Assistant Government Entomologist, Kenya Colony.

The first record of red scale in Kenya Colony in the file of the Entomological
Laboratory is for 1914, on citrus. It is known that the scale has been brought into
the country in at least one consignment of citrus, which, being accompanied by a
certificate from the country of origin that the plants were free from insect pests, was
allowed to enter the country under the Plant Import Regulations then in force, without
being inspected. It seems, therefore, likely that red scale is not indigenous to the
country, but an introduced pest.

In this belief, an attempt at eradication was made, in the hope that the insect might
be prevented from becoming established in the country, and the citrus crop thus
saved from a costly enemy. This attempt was also undertaken because of the possi-
bility that red scale might attack coffee, a very much more important crop in the
country than citrus. This was recognised as a danger for two reasons : first, because
red scale in other countries has a very wide range of food-plants ; and secondly,
because quite a number of scales in Kenya Colony attack both citrus and coffee.
These are: Icerya purchasi, Mask., Saissctia hemisphaerica, Targ., Saissetia nigra,
Niet., Coccus hesperidum, 1.., Ceroplastes ceriferus, And., and Selenaspidus articulatus,
Morgan.

Accordingly, the importation of citrus into the country was prohibited. At the
same time, owners of citrus attacked by red scale were advised to destroy such trees,
which were replaced free from the Government Farm, Kabete. Compulsory powers
were not sought, but whenever planters were asked to destroy infected trees under
this scheme, they consented to do so.

Commencing in 1917, inspections of citrus orchards and nurseries in search of red
scale were made by the staff of the Division of Entomology and by the late Mr. J. J.
Adams. A circular letter was sent out to citrus owners, accompanied by a specimen
of red scale, asking them to make a search for the insect on their trees and to report
the result. Sometimes it was found that other insects, such as Coccus hesperidum
or Selenaspidus articulatus, or the citrus Psyllid, Tvioza sp., were mistaken for red
scale, so owners who reported the presence of this insect were asked to send a specimen
from their trees.

By these means it was found that red scale was widely distributed in the country
and that large numbers of trees were attacked. In some cases, citrus not being a
profitable crop, little care has been taken of the trees and the scale has been allowed
to multiply unchecked, so that in some neglected orchards great damage has been
done by it.

In addition to citrus the plants now known to be hosts of red scale in this
country are roses, apple, plum and sisal.

Fortunately, the fears that red scale might attack coffee have not so far been
realised, and the following evidence, obtained both in the field and in the laboratory,
is encouraging :—

(1) Red scale has never been observed or reported on coffee.

(2) A field of coffee, adjacent to some citrus trees, very badly infested with red
scale, has been kept under observation. The citrus is very close to the
outside row of coffee, the branches of the two often being in contact. All
that has been found on the coffee, are objects that appeared to be red
scale larvae, which have secreted the white covering just after settling
(this the larvae have also done when put on coffee in the laboratory) and
older individuals that were dead and had in all probability been washed
off the citrus on to the coffee.

104 F. W. DRY.

(3) In experiments in the laboratory many hundreds of red scale larvae have
been placed on the leaves of coffee plants. Many of them secreted their
white covering, but none made any further progress. Larvae similarly
placed on citrus as controls lived and attained maturity and themselves
produced offspring.

On the citrus plants, to which reference has been just made, the life-history of the
red scale was worked out, the larvae having been placed on the plants in the middle
of November 1917. The times when the various stages in the life-history were first
teached were as follows :—

After the first day most of the larvae had secreted a covering.

After eight days the “ pimple ”’ was visible in the centre of the scale.

The first moult took place after 18 days.

The second moult of the female took place after 50 days.

The first male emerged on the 65th day.

The first larvae of the next generation were observed after 110 days.

The average daily temperatures during this life-cycle were : minimum, 54:5° F. ;
maximum, 80:5° F.

Unfortunately the eradication of red scale has proved impracticable, but we may
hope that coffee is immune.

Citrus is not at present an important crop in the country, though a great many
people have a small number of trees. At local prices fumigation is impracticable,
so that spraying is the control measure recommended.

The West Indian red scale, Selenaspidus articulatus, Morgan, has several times been
mistaken for the common red scale by citrus owners. It occurs on both citrus and
coffee, and has been found in widely separated parts of the country, but has not been
known to do serious harm ; often only a single individual will be found on one leaf.
In one citrus orchard in the Songhor district, a small number of trees could be described
as badly infested. The fruit was attacked as well as the leaves, hundreds of
individuals being found on a single orange, which was thus rendered quite unsightly.

A citrus plant was infected in the laboratory, at the same time as those on which
the red scale life-history was worked out. Larvae of the next generation were first
observed after 120 days.

105

COLLECTIONS RECEIVED,

The following collections were received by the Imperial Bureau of Entomology
between Ist January and 31st March, 1921, and the thanks of the Managing Committee
are tendered to the contributors for their kind assistance :—

Mr. E. BALrLarpD, Government Entomologist :—3 Tachinidae, 30 Coleoptera,
100 Thysanoptera, 60 Rhynchota, and a collection of Nematode worms suspected of
damaging crops; from Madras.

Mr. G. E. Bopxkin, Government Economic Biologist :—35 Culicidae, 5 Tabanidae,
15 other Diptera, 14 Hymenoptera, 141 Coleoptera, 6 Moths, 50 Isoptera, 2 species
of Coccidae, 41 other Rhynchota, 2 Orthoptera, 5 Odonata, 44 Spiders, 4 Scorpions,
2 Centipedes, 5 Millipedes, 2 Worms, and 10 insects attacked by Entomogenous
fungi; from British Guiana.

Mr. Harotp E. Box :—8 Tabanidae, 13 other Diptera, and 4 species of Coccidae ;
from Argentina.

Mr. P. A. Buxton :—30 Rhynchota ; from Mesopotamia.

Dr. G. D. H. CARPENTER :—10 Tabanidae, 5 other Diptera, 9 Hymenoptera,
198 Coleoptera, 34 Rhynchota, 26 Orthoptera, and 2 Stone-flies ; from Uganda.

Mr. J. B. CorporaaL, Entomologist, Algemeen Proefstation, Medan :—-8 Cur-
culionidae and 23 Rhynchota ; from Sumatra.

Mr. M. T. Dawe :—1 Tabanus, 15 Glossina, 60 Coleoptera, 9 Lepidoptera, and
1 species of Coccidae; from Gambia; and 8 Tabanidae and 111 Ticks; from
Colombia.

Mr. D. pD’EMMEREZ DE CHARMOY :—1 Tube of Cecidomyiidae, 6 Lepidoptera,
1 species of Coccidae, and 300 Aphididae; from Mauritius.

Dr. EustacE W. FERGUSON :—37 Tabanidae; from Australia.

Mr. T. BAINBRIGGE FLETCHER, Imperial Entomologist :—30 Anoplura ; from India.

_Mr. C. C. GowpEy, Government Entomologist :—220 Ants, 32 Chalcids, 3 other
Hymenoptera, 10 Coleoptera, 7 Lepidoptera, 50 Thysanoptera, 7 species of Coccidae,
3 other Rhynchota, 1 Dragon-fly, and 45 Mites ; from Jamaica.

Mr. E. HARGREAVES :—70 Sheep “ Ticks,” 13 Lepidoptera, and 2 Orthoptera ;
from the United Kingdom : and 2 Mutillidae, 115 Coleoptera, and 5 Rhynchota ;
from North America.

Mr. H. HARGREAVES, Government Entomologist :—10 Hymenoptera, 123 Coleop-
tera, and 49 Rhynchota ; from Uganda.

Mr. G. F. Hitt, Entomologist, Australian Institute of Tropical Medicine :—191
Culicidae, 59 Tabanidae, 57 other Diptera, 6 Ants, 5 Chalcids, 16 Coleoptera,
12 Isoptera, 5 species of Coccidae, and 4 other Rhynchota ; from Australia.

Dr. WALTHER Horn :—4 specimens of Chrysops costata, F.; from Cuba.

Mr. M. ArzaL Husain, Government Entomologist :—54 Diptera and 33 Chalcids ;
from the Punjab.

Mr. J. C. Hutson, Government Entomologist :—387 Parasitic Hymenoptera, 39
Ticks, and 6 other Arachnida ; from Ceylon.

Mr. F. P. Jepson, Assistant Government Entomologist :—38 Coleoptera ; from
Ceylon.

Mr. L. Lewron-Brain, Director of Agriculture :—5 Diptera, 91 Coleoptera, 24
Lepidoptera, and 14 Orthoptera ; from the Federated Malay States.

Major W. F. M. LoucHnan, R.A.M.C.: 4 Tubes of Ceratopogoninae; from
Jamaica and British Honduras.

(2416) H

106 COLLECTIONS RECEIVED.

Mesopotamia, Director of Agriculture :—40 Diptera, 13 Hymenoptera, 37 Coleop-
tera, 28 Lepidoptera, and 46 Rhynchota ; from Mesopotamia.

Mr. F. Murr :—14 Homoptera ; from Samoa, etc.

Mr. H. K. Munro :—18 Coleoptera ; from South Africa.

Prof. G. H. F. Nutra, F.R.S. :—2 Tabanidae, 22 Stomoxys, 2 Hippoboscidae,
and 108 other Diptera ; from Mesopotamia.

The Rev. Father J. A. O’NEtz, S.J. :—87 Coleoptera and 7 Moths ; from Southern
Rhodesia.

Dr. L. PértNcueEy, Director of the S. African Museum :—4 Weevils; from Cape
Colony.

Philippine Islands, Director of Agriculture :—8 Orthoptera ; from the Philippines.

Mr. A. W. J. Pomeroy, Government Entomologist :—14 Diptera, 400 Chalcids,
30 other Hymenoptera, and 28 Lepidoptera ; from Nigeria.

Pretoria, Division of Entomology :—84 Orthoptera ; from South Africa.

Mr. A. H. Rircute :—100 Ants, 3 Chalcids, 152 Coleoptera, 13 species of Coccidae,
and 200 Aphididae; from Jamaica.

Senhor A. F. pE SkaBra :—18 Orthoptera and 2 Odonata ; from San Thome.

Dr. B. P. Uvarov :—28 Hymenoptera, 34 Coleoptera, and 4 Rhynchota; from
Transcaucasia.

Mr. Ropert VEITcH :—29 Diptera, 12 Chalcids, 16 other Hymenoptera, 118
Coleoptera, 50 Lepidoptera, and 30 Rhynchota ; from Fiji.

Sir Francis Watts, K.C.M.G., Imperial Commissioner of Agriculture :—400
Chalcids, 50 Coleoptera, 37 Lepidoptera, 4 species of Coccidae, Aleurodidae and
Aphididae, and 2 other Rhynchota; from the British West Indies.

Wellcome Bureau of Scientific Research :—-26 Meloid beetles ; from Peru.
Mr. C. B. Witirams :—4 Coleoptera and 13 Rhynchota ; from Trinidad.

SEPTEMBER, 1921.

VOL. XII. Part 2.—pp. 107-204.

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107

A CONTRIBUTION TO KNOWLEDGE OF THE BLOOD-SUCKING
DIPTERA OF PALESTINE, OTHER THAN TABANIDAE.*

By Major FE. E. Austen, D.S.0O.
(Plate DV)

As in the case of the TABANIDAE, which have already been described,* the
material upon which the following paper is based was collected by the author
during the Palestine Campaign of 1917-18.

All the specimens, including types of new species, are in the British Museum

(Natural History).

Family CHIRONOMIDAE.
Sub-family CERATOPOGONINAE.
Genus Leptoconops, Skuse.
Syn. Tersesthes, Townsend.

_Leptoconops kerteszi, Kieffer.t

In abundance near Wadi Ghuzze, on Cairo road about 5 miles S.-W. of Gaza,
on afternoon of 14.v.1917; settling in numbers on the faces of the writer and a

companion.

L. kertészi, Kieffer, originally described (Ann. Mus. Nat. Hung., vi, pp. 576-577,
1908) from material taken at Cairo, and subsequently recorded by. Kieffer (op. cit.,
xvi, p. 34, 1918) as occurring in Tunisia, is already represented in the National
Collection by a series of specimens from Ouargla, Algeria (Dr. E. Hartert), March 1912,
bearing the following field-note by the collector :—‘* Exceedingly numerous in some
of the oases south of Biskra, and very troublesome to mules.”’ It may be added
that at Bir el-Abd, Northern Sinai (50 miles E. of Kantara), 9.xii.1916, the writer
met with two females of what appeared to be this species on the margin of a small
salt lake, and was bitten by one of them on the arm at midday.

In life the dorsum of the abdomen of this little midge shows a double, longi-
tudinal series of admedian, dark brown blotches, separated by neutral grey?, triangular
interspaces; the venter is whitish ; the wings, the surface of which is apparently
bare, are uniformly milk-white, except that in each wing the fused ends of the first
and third longitudinal veins are expanded to form a kind of stigma, which is large
and very conspicuous, and of a striking orange colour ; the halteres are pale buff.

The suggestion by Kieffer (op. cit., vi, p.577) that Tersesthes, Townsend (founded
for a species which attacks horses at fairly high altitudes—5,700—7,000 ft—in New
Mexico) is probably identical with Leptoconops, Skuse, is undoubtedly correct.

* For Tabanidae, cf. the author’s paper “‘ A Contribution to Knowledge of the Tabanidae
of Palestine’”’: Bull. Ent. Res., x, pt. 3, pp. 277-321, figs. 1-18 (April 1920).

+ In 1918 this species was selected by Kieffer as the type of a new genus, which he briefly
characterised (Ann. Mus. Nat. Hung., xvi, p. 135, 1918) under the name Holoconops, relying
upon the number of joints in the antenna of the 2 to justify a generic distinction. As was
recently shown, however, by Mr. H. F. Carter in his admirable ‘ Revision of the Genus Lepto-
conops, Skuse’’ (Bull. Ent. Res., xii, pt. 1, pp. 1-28, June 1921), it is impossible to accord to
Holoconops anything more than subgeneric rank.

+ For names and illustrations of colours used for descriptive purposes in the present paper,
see Ridgway, “‘ Color Standards and Color Nomenclature’ (Washington, D.C. Published by the

Author, 1912).

(3442) 8/170 1000 8/21 Harrow G75. I

108 MAJOR E. E. AUSTEN.
Genus Culicoides, Latr.

Synoptic Table.

The eight species described or recorded below are mutually distinguishable
as follows :—

1 (4) Wings entirely devoid of markings.
3

2 (3) Mesonotum and crown of head olivaceous black .. .. vttreipennts, sp. Nn.
3 (2) Mesonotum and crown of head grey (greyish olive) .. puripennts, sp. n.
4 (1) Wings with markings.

5 (8) Pale markings oneach wing (other than any that there may be at the extreme

base) limited to two more or less conspicuous spots or flecks on or close
to costal border.
6 (7) Pale wing-markings on costal border clearly defined and conspicuous ;
mesonotum cinnamon-coloured .. 5 CentOriuS, Sp. aus
7 (6) Pale wing-markings on or near costal border i inconspicuous, and from certain
angles visible with difficulty ; mesonotum dark mummy-brown
odiatus, sp. Nn.
8 (5) Pale markings on each wing (other than those at the extreme base) not
limited to two spots or flecks, but much more numerous and extensive.
9 (10) Wings each with three dark blotches on costal border, of which at least
the two more distal are (as seen cae a light Pee) conspicuously
darker than those elsewhere ee .. newsteadt, sp. n.
. Wings not so marked ; darker blotches on ‘costal border (as seen against a
light background) not conspicuously deeper in tint than those elsewhere.
12) Mesonotum unicolorous ar a 3 ve .. guttularts, Kieffer.
12 (11) Mesonotum not unicolorous.
14) Mesonotum with light grey markings on a dark brown ground; wings
with pale spots at their distal extremities directly in contact with the
wing-margin, no dark spot in contact with anterior transverse vein
odibilis, sp. n.
14 (13) Mesonotum speckled; wings in each case with distal pale spot between
rami of fourth longitudinal vein not directly in contact with wing-margin,
a small but conspicuous dark spot in contact with anterior transverse vein
circumscriptus, Kieffer.

Culicoides vitreipennis, sp. n.

°.—Length (one dried specimen), from anterior margin of thorax to posterior
extremity of abdomen, 1 mm.; length of wing, 1:25 mm., greatest breadth of
wing, 0:6 mm.

Wings hyaline, with a milky appearance but entirely devoid of markings ; crown
of head and dorsum of thorax olivaceous black; legs pale.

Head: vertex sparsely clothed with short, ochreous hairs, space between upper
lobes of eyes with longer ochreous hairs; face and proboscis dark brown ; inner
margins of upper lobes of eves moderately wide apart ; palpi mummy-brown, third
segment oval, moderately swollen; second segment of antennae (torus) mummy-
brown, relatively large (larger and darker than in the following species), flagellum
drab, tinged with brownish towards distal extremity, clothed with yellowish hair,
first five or six segments of flagellum more or less spherical in shape, more truncate
(more spherical) than in the following species. Thorax entirely without markings,
mesonotum sparsely clothed with short, ochreous hairs ; scuwtellum agreeing in colora-
tion with remainder of dorsum, and bearing one lateral bristle on each side, also
(apparently) one central bristle, as well as several short hairs. Abdomen (in dried
condition} blackish brown, clothed at distal extremity with yellowish hairs. Wings

THE BLOOD-SUCKING DIPTERA OF PALESTINE. 109

clothed (somewhat less extensively than in following species) with fine, pale, decum-
bent hairs (macrotrichia), wing-fringe likewise pale; costa as far as end of third
vein, first and third longitudinal veins, base of fourth vein as far as anterior
transverse vein and anterior transverse vein itself pale cream-buff, otherwise all
veins colourless ; distal extremity of third longitudinal vein curved round (not bent at
an angle) to meet costa, anterior transverse vein if anything slightly shorter than in
following species; both radial cells distinct, but not quite so large as in latter.
Halteres ivory-yellow, stalks brownish at base. Legs cream-buff, femora, except
at distal extremities, and tibiae, except at their bases, tinged with sepia, extreme
tips of femora and extreme bases of tibiae dark brown, extreme tips of hind tibiae
mummy-brown on inner side; hair on legs pale yellowish, longer hairs on outer
surface of hind tibiae inconspicuous.

Near Jerisheh, 5 miles N.-E. of Jaffa, 1-8.v.1918, in author’s tent at night,
on lining, above lighted lamp.

The species just described is distinguishable from the European C. pumilus,
Winn., which it resembles in size and in the coloration of the body, by the much paler
wings and legs, and by the decumbent hairs (macrotrichia) on the wings being pale
instead of dusky.

Culicoides puripennis, sp. n.

°.—Length (one dried specimen), from anterior margin of thorax to posterior
extremity of abdomen, 1-25 mm.; length of wing, 1-4 mm., greatest breadth of
wing, 0-6 mm.

Wings hyaline, somewhat milky, entirely devoid of markings ; crown of head and
dorsum of thorax deep greyish olive; legs pale.

Head: vertex clothed with pale yellowish hairs, face greyish sepia-coloured ;
proboscis mummy-brown ; inner margins of upper lobes of eves closely approximate ;
palpi sepia-coloured, third segment but slightly swollen, first three segments
clothed mainly with dark brown or blackish hair, last two segments clothed
with ochreous hair; second segment of antennae (torus) light sepia-coloured, paler
and also somewhat smaller than in foregoing species, flagellum drab, clothed with
yellowish hair, five segments following torus bluntly ovoid. Thorax entirely without
markings, mesonotum clothed with pale yellowish hairs ; scutellum agreeing in
coloration with remainder of dorsum, and apparently bearing two central bristles
and on each side two lateral bristles, as well as several short hairs. Abdomen (in
dried condition) clove-brown, sparsely clothed with pale hairs. Wangs: greater
part of surface, except base and costal, basal and radial cells, clothed with fine, pale,
decumbent hairs (macrotrichia), which at and towards distal extremity of each
wing are shorter and more closely set ; wing-fringe pale; costa as far as end of
third vein, first and third longitudinal veins, base of fourth vein as far as anterior
transverse vein and anterior transverse vein itself cream-buff (distal half of third
longitudinal vein and portion of costa immediately above it somewhat darker),
otherwise all veins colourless; distal extremity of third longitudinal vein bent up
at an obtuse angle to meet costa, anterior transverse vein fairly long ; both radial
cells well developed, the first about half as long again as the second. Halteres :
knobs straw-yellow, stalks slightly darker. Legs cream-buff or pale cinnamon-buff,
distal extremities of hind femora and hind tibiae sepia-coloured, extreme tips of
middle femora and tibiae tinged with brown, first joint of hind tarsi somewhat
infuscated ; hair on legs pale, inconspicuous.

Deiran (Rechoboth), Jaffa district, 7 miles S.-W. of Ludd, 12.iv.1918, in room
at night.
(3442) 12

110 MAJOR E. E. AUSTEN.

From the foregoing species C. puripennis is distinguishable, inter alia, by the
coloration of the dorsum of the thorax ; by the second segment of the antenna being
smaller and paler and the following five segments somewhat more elongate ; and by
the course (angulate instead of rounded) followed by the distal extremity of the
third longitudinal vein.

The species just described is also allied to the European C. albicans, Winn., from
which it may be distinguished owing to the front and middle femora and tibiae not
being conspicuously tipped with blackish brown or black, and also (if Winnertz’s
figure—Linnaea Entomologica, vi, Taf. vi, fig. 35) (1852) is to be relied upon) by
the less abruptly turned up distal extremity of the third longitudinal vein.

Culicoides tentorius, sp. n. (PI. iv, fig. 1).

g.—Length (3 dried specimens), from anterior margin of thorax to posterior
extremity of abdomen, 1-2 to 1-4 mm. ; length of wing, 1-5 to 1-75 mm. ; greatest
breadth of wing, 0-6 mm.

9.—Length (9 dried specimens), from anterior margin of thorax to posterior
extremity of abdomen, | to 1-5 mm. ; length of wing, 1:4 to 1-6 mm., greatest
breadth of wing, 0°75 mm.

_ Dorsum of thorax (in dried specimens) cinnamon-drab_ (3, and sometimes 9), or
cinnamon-coloured (9) ; dorsum of abdomen (in dried specimens) clove-brown or warm
sepia-coloured ; wings mouse-grey (3), or sepia-coloured or dusky-drab (Q), an obliquely
elongate area at base pale, otherwise in both sexes the only light markings ave situate

. . . . = ° a
on costal border in shape of a pair of conspicuous rvory-yellow spots, of which that nearer
the base as the larger (Pl. iv, fie. 1) ground-colour of legs cream-buff or cinnamon-buff,
hind tibiae in both sexes fringed posteriorly with a row of long hairs.

Fig. 1. Culicoides tentorius, sp.n.; palpusof 9.

Head fawn-coloured (face cinnamon-buff), vertex infuscated (blackish-brown or
deep mouse-grey) in g and in both sexes clothed with curved, glistening, yellowish
hairs ; inner margins of upper lobes of eyes in 2 very narrowly separated ; proboscis
and palpi cinnamon-buff in 3, sepia-coloured or dark brown in 9, palpi in clothed
with brownish hair, third segment moderately swollen towards distal extremity
(fig. 1) ; antennae in both sexes cream-buff or light cinnamon-buff (distal segments
sometimes darker), fourth to tenth segments in 9 moderately elongate (fig. 2), hair
on antennae, including antennal plume in 4, glistening yellowish (light ochreous or
ochraceous-buff). Thorax: dorsum without markings in either sex, sparsely clothed
with glistening ochraceous-buff hairs ; scutellum agreeing in coloration with remainder
of dorsum, and bearing in both sexes two central and two lateral bristles, as also
several short hairs. Abdomen in both sexes clothed with pale yellowish hair.
Hypopygium of 3 (fig. 3): ninth sternite deeply notched in middle line ; posterior

THE BLOOD-SUCKING DIPTERA OF PALESTINE, Dit

margin of ninth tergite also deeply notched, and with relatively broad finger-like
extensions ; lobe-like processes of lower surface of projecting portion of ninth tergite
situate some distance in front of posterior margin ; forceps of usual form, side-pieces
each with two slender, sub-dorsal processes on inner side of proximal extremity ;
proximal portion of each harpfe with a strongly chitinised ventral process, at right
angles to distal portion, distal portions of harpes noticeably broad, and each tapering

y pte
Wg (SE NS
YES YAS
GEA.
mec S

£.6.Q,
Fig. 2. Culicoides tentorius, sp.n.; antenna of Q.

to a point posteriorly ; aedoeagus Y-shaped or lyrate, with broad, well chitinised
stem, terminating bluntly behind, and with strongly chitinised limbs. Wangs
(Pl. iv, fig. 1) agreeing in both sexes as regards markings ; proximal ivory- yellow
spot on costa surrounding anterior transverse vein, varying somewhat in size and
outline in different individuals, but with its lower extremity reaching fold which
traverses fork of fourth longitudinal vein ; proximal boundary of distal spot formed

Fig. 3. Culicoides tentorius, sp. n.; male hypopygium, ventral view (greatly enlarged).

by or scarcely extending beyond terminal upturned portion of third longitudinal
vein ; in both sexes portions of costa and of first and third longitudinal veins between
spots conspicuously darker than anything else in wing ; a larger or smaller area at base
of wing pale, portions of veins included in this area, as also anterior transverse vein,
upturned terminal portion of third longitudinal vein, and portions of longitudinal
veins in proximal costal spot light buff or cream-buff, veins elsewhere, except as already

112 MAJOR E. E. AUSTEN.

stated, sepia-coloured ; in 9, decumbent hairs absent from base of wing, including
basal cell and costal border as far as end of first longitudinal vein, remainder of
wing well clothed with these hairs, which are especially close together on costal
border beyond distal costal spot. Halteres in both sexes cream-coloured or cream-
buff. Legs: femora and tibiae, at least of hind legs, usually darker (tinged with
mummy-brown or sepia) except at base and tip, so that these joints have a pale band
at each extremity, extreme tips of femora and tibiae, at least of hind legs, often
mummy-brown.

Near Jerisheh, 5 miles N.-E. of Jaffa, 26.iv.—8.v.1918: type of g and 4 ¢
para-types, type of 2 and 10 @ para-types, in author’s tent at night, on lining,
above lighted lamp. Not observed biting.

Although in general appearance, including coloration and wing-markings, closely
resembling the East and West African Culicoides ( Johannseniella) fulvithorax,
Austen, the species just described is distinguishable therefrom, inter alia, by its
generally larger size, by the less elongate shape of the fourth to the tenth segments
inclusive of the 9 antenna, and by the fact that in the wing the distal pale spot on
the costa, instead of surrounding the terminal portion of the third longitudinal
vein, is only in contact with the upturned part of its extreme tip.

From the N. Italian Culicoides susae, Kieff., C. tentorius may be distinguished
by its larger size, by the fourth to the ninth segments inclusive of the 2 antenna not
being globular, by the halteres being cream-buff or cream-coloured instead of white,
and by the presence of a row of long hairs on the extensor surface of the hind tibiae.

Culicoides odiatus, sp. n.

°.—Length (2 dried specimens), from anterior margin of thorax to posterior
extremity of abdomen, 1-4 to 1-5 mm.; length of wing, 1-4 to 1-5 mm., greatest
breadth of wing, 0:6 to 0-8 mm.

Dusky species, with very hairy wings, which are almost entirely devoid of markings.—
Head and thorax dark mummy-brown (vertex darker), without spots or other markings ;
abdomen (in dried specimens) russet-brown or russet; wings with markings in each
case confined to a pair of small, faint, ill-defined and inconspicuous maculae on or near
costal border.

Head clothed above with brownish hair; inner margins of upper lobes of eyes
almost in contact below ; proboscis cinnamon-brown ; palpi dark mummy-brown,
clothed with dark brown hair, third segment strongly swollen, expansion com-
mencing immediately beyond base ; antennae light sepia-coloured, agreeing closely
with those of C. tentorius, Austen -(cf. fig 2, p. 111) as regards shape of segments,
and clothed with yellowish (ochraceous-buff) hairs. Thorax: mesonotum clothed
with ochraceous-buff hairs and with dark brown bristles; scztellum bearing, in
addition to several short hairs, apparently four central bristles, as also on each side
two lateral bristles. Abdomen sparsely clothed with pale hairs, except on sides
and at distal extremity, where a portion at least of the hairs are dark brownish.
Wings : with exception of basal and costal cells, practically entire surface clothed
with long, closely set, decumbent hairs, which, especially in region beyond level of
end of third longitudinal vein, where they are thickest, largely overlap one another ;
of the two faint pale maculae in each wing, that nearer the base surrounds the anterior
transverse vein and extends only indistinctly to the costa, the other is situate on and
adjacent to the costa at the end of the third longitudinal vein, the extreme tip of
which it includes ; second radial cell much broader than the other, which is practically
obliterated and indistinguishable. Halteres cream-buff, distal extremities of stalks
greyish. Legs: femora light sepia-coloured (hind pair darker), in each case with a
faintly marked pale band (less conspicuous in that of hind pair) before extreme tip,

THE BLOOD-SUCKING DIPTERA OF PALESTINE. 1S

which is infuscated; tibiae drab-coloured or brownish drab, their extreme tips
infuscated ; tarsi light ochraceous-buff ; tibiae clothed with brownish or yellowish
hair, a series of long hairs on extensor surface of hind pair ; upper surface of hind
tarsi clothed with fairly long, pale hair.

Near Jerisheh, 5 miles N.-E. of Jaffa, 29.iv.—8.v.1918: type and one _ para-
type, in author’s tent at night, on lining, above lighted lamp.

The species described above is allied to the foregoing (C. tentorius, Austen), but,
in the 9 sex at any rate, is distinguishable znter alia by the more swollen third joint
of the palpi, by the much darker colour of the dorsum of the thorax, by the wings
being much more hairy, and by the two pale spots on the costal border being only
faintly indicated and much less developed.

Culicoides newsteadi, sp. n. (Pl. iv, fig. 3).

Q.—Length (3 dried specimens), from anterior margin of thorax to posterior
extremity of abdomen, 1-2 mm.; length of wing, 1-3 mm., greatest breadth of
wing, 0-6 mm.

Allied to the European C. pulicaris, L., and agreeing therewith in wing-markings
in case of specimens in which pattern shown in Pl. iv, fig. 3 is somewhat reduced,
but, in 2 sex at any rate, distinguishable inter alia by much smaller size, and by presence
of a pale band, sharply defined in case of fully-coloured specimens when viewed against
a dark background, at distal extremity as well as at base of hind tibiae.

Head: vertex sparsely clothed with curved, yellowish hairs; inner margins of
upper lobes of eyes in contact or separated by an exceedingly narrow interval ;
proboscis dark brown; palpi sepia-coloured, clothed partly with brownish, partly
with yellowish hair, third segment strongly swollen; antennae light sepia-coloured
or light mummy-brown, third to tenth segments inclusive generally paler (cream-
coloured), hair on antennae yellowish. Thorax: dorsum clothed with shining
ochraceous-buff hairs, ground-colour (in dried specimens) olive-grey, with a mummy-
brown area on each side anteriorly, or light greyish olive, with anteriorly a narrow,
sepia-coloured, longitudinal streak in middle line, becoming obsolete towards hind
margin, and on each side, between it and lateral border, a broader and longer, curved,
longitudinal stripe of same colour; scwtellwm agreeing in ground-colour with
remainder of dorsum, and bearing two central and two lateral bristles, as also three
short hairs between each central and corresponding lateral bristle: Abdomen
sparsely clothed with pale (cream- or cream-buff-coloured) hairs. Wings: in speci-
mens with fully-developed wing-markings, latter are as shown in PI. iv, fig. 3, the
three dark blotches on costal border dark mouse-grey and very conspicuous, remaining
dark markings mouse-grey ; in many specimens, however, mouse-grey markings
between costal border and hind margin are much reduced in extent, taking form of
partly discontinuous and isolated spots and flecks; greater part of distal half of
wing-surface, as well as of hind border, fairly thickly clothed with decumbent hairs.
Halteres : stalks cream-coloured, knobs ivory-yellow. Legs sepia-coloured or light
sepia-coloured, tarsi, middle and hind knees, and a band at each extremity of hind
tibiae paler (pinkish-buff or pale pinkish-buff—in well-coloured specimens pale bands
on hind tibiae are cinnamon-buff) ; hair on legs pale, hind tibiae on outer surface
with a row of long hairs.

Near Jerisheh, 5 miles N.-E. of Jaffa, 26.iv—8.v.1918: type and 5 para-types,
in author’s tent at night, on lining, above lighted lamp: dedicated, as a trifling token
of sincere regard, to Robert Newstead, F.R.S., Dutton Memorial Professor of
Entomology, Liverpool School of Tropical Medicine. Although not actually taken
in flagrante delicto, there can be no doubt that this species is a blood-sucker, since
in the case of the type fresh blood was observed in the abdomen at the time of capture.

114 MAJOR E. E. AUSTEN.

A 3 Culicoides taken, with specimens of four other species of the same genus,
at Sheikh Zowaiid, N. Sinai, 14.11.1917, in the author’s tent at night, and almost
certainly belonging to the species just described, measures (in the dried condition)
1 6 mm. in length from the anterior margin of the thorax to the posterior extremity
of the abdomen. In the genitalia, the side-pieces (basal portions of forceps) are
considerably less swollen than in the case of C. pulicaris, L., J, so that the disparity
in size between these and the claspers is not so great. As regards wing-markings,
with the exception of the distal costal blotch, which is undiminished, all the dark
markings are much reduced, those other than on the costal border appearing as
some eight or nine faint and almost completely isolated flecks.

Culicoides guttularis, Iieff.
Culicoides guttularis, Kieffer, Ann. Mus. Nat. Hung., xvii, p. 45 (1919).

Three 99, near Jerisheh, 5 miles N.-E. of Jaffa, 29.iv.—8.v.1918, in author’s
tent at night, on lining, above lighted lamp. Not observed biting, but one of the
three specimens brought back has its abdomen distended, apparently with blood.

C. guitularis, the type of which was taken in Hungary (Budapest), also occurs in
Great Britain, and has been taken in Middlesex, Herts, Huntingdonshire and the
Isle of Arran. The specimens obtained in Palestine differ from the typical form,
as described by Kieffer, and agree with British examples, in having a dark transverse
mark across the centre of the axillary cell, the distal extremity of which, in contact
with the posterior branch of the fifth longitudinal vein, is also infuscated.

Culicoides odibilis, sp. n. (Pl. iv, fig. 2).

g.—Length (1 dried specimen), from anterior margin of thorax to posterior
extremity of abdomen, 1:25 mm.; length of wing, 1-5 mm., greatest breadth of
wing, 0:6 mm.

Antennal plume cream-buff (looking brownish in certain aspects, or when the hairs
are matted together) ; dorsum of thorax dark brown, with conspicuous light grey mark-
ings ; wings infuscated and iridescent, with sharply defined milk-white or cream-coloured
spots, as shown in PI. iv, fig. 2.

Head: proboscis and palpi mummy-brown, clothed with dusky hairs; torus of
antenna blackish brown, third to twelfth segments inclusive ivory-yellow or almost
colourless, last three segments sepia-coloured in certain lights, clothed with dusky
hairs mixed with some pale hairs. Thorax dark mummy-brown, mesonotum
sparsely clothed with glistening yellowish hairs, and with ground-colour varied by
conspicuous and sharply defined light neutral grey markings, chief among which are
a sinuous mark, shaped something like a note of interrogation (?), embracing each
anterior slit-like depression, a small spot behind each of these marks, a transverse
row of four transversely elongate spots across the middle, and a broad, somewhat
trident-shaped mark on hind border ; base and sides of scutellum light neutral grey.

Abdomen dark neutral grey (claspers paler—light brownish olive), clothed
with dusky hair, paler (yellowish) towards the tips. Hypopygium : ninth
sternite deeply emarginate; posterior margin of ninth tergite not noticeably
notched in middle line, finger-like extensions narrower than in C. guttularis,
Kieff.; lobe-like processes of lower surface of projecting portion of ninth
tergite situate some distance in front of posterior margin ; forceps of usual form,
but claspers less elongate than in C. guttularis, and their distal extremities
not swollen as in the latter species ; harpes broad, with distal extremity of each
attenuate and elongate; limbs of aedoeagus strongly chitinised and forming
a Y-shaped or lyrate rather than a U-shaped figure, as in C. guttularis, stem of
aedoeagus moderately broad, apparently rather short, fairly well chitinised and
ending bluntly behind. Wings (Pl. iv, fig. 2) deep mouse-grey, with a strongly

THE BLOOD-SUCKING DIPTERA OF PALESTINE. 115.

developed purplish iridescent sheen, and with sharply defined, spot-like, milk-white
or cream-coloured markings, chiefly along distal and posterior borders, as shown in
figure ; hind border of axillary cell to level of axillary angle, and greater part of
distal half of each wing fairly thickly clothed with decumbent hairs, which as usual
are especially close together on distal third, particularly in area beyond distal costal
spot. Halteres: knobs straw-yellow, stalks cream-coloured. Legs: femora and
tibiae light sepia-coloured (hind femora somewhat darker), knees (tips of femora and
extreme bases of tibiae) and tips of hind tibiae mummy-brown ; tarsi, extreme bases
of all femora, a ring immediately before tips of front and middle femora, and a similar
ring immediately beyond bases of all tibiae and before tips of hind tibiae cream-
coloured ; hind femora with a faint indication of a narrow pale ring before tips ;
hind tibiae with a row of long, dusky hairs on extensor surface, hair on tarsi and on
flexor surface of hind tibiae pale.

Near Jerisheh, 5 miles N.-E. of Jaffa, 26.iv.1918; in author’s tent at night,
on lining, above lighted lamp.

Culicoides odtbilts is allied to the foregoing species (C. guttularis, Kieff.), from which
however it is readily distinguishable by the neutral grey markings on the dorsum
of the thorax, by the differences in the § hypopygium detailed above, and by the
much more sharply defined wing-markings, in which the pale spots are considerably
less extensive.

Culicoides circumscriptus, Kieff.
Culicoides circumscriptus, Kieffer, Ann. Mus. Nat. Hung., xvi, p. 49, fig. 15 (1918).

One 9, near Jerisheh, 5 miles N.-E. of Jaffa, 29.iv.1918, in author’s tent at
night, on lining, above lighted lamp.

The specimen referred to agrees on the whole very well with Kieffer’s description
of the species, the type of which was obtained in Tunis. The inner margins of the
upper lobes of the eyes, though separated above by a space, the width of which is
approximately equal to that of the flagellum of the antenna, are closely approximate
below. In the much bespotted wings, practically the entire surface of which is thickly
clothed with decumbent hairs, the dark fleck (alluded to by Kieffer) enclosed in the
proximal pale costal blotch is situate immediately beyond, and in contact with, the
anterior transverse vein, occupying the angle formed by the latter and the praefurcal
portion of the fourth longitudinal vein; a small pale fleck inside the fork of the
fourth vein, close to its base, is not mentioned by Kieffer.

Note——In an addendum to a short paper by E. Brunetti on “‘ Some Noxious
Diptera from Galilee”’ (Journ. Asiatic Soc. Bengal, New Ser., ix, pp. 43-35, 1913),
Dr. N. Annandale writes (loc. cit., p. 45) :—‘‘ Another irritating blood-sucker common
at Tiberias in October, though much less so than Ph. papatast, is a minute Chiro-
nomid of the sub-family Ceratopogoninae. Like Phlebotomus it is nocturnal in its
habits.” On p. 370 of Vol. x of the same journal (1914), in a note to Kieffer’s
description of Trichotanypus tiberiadis, Wieff., Dr. Annandale remarks :—“ This is
the species I referred to in a note on a former paper (J..A. S.B. (n.s.), ix, p. 45, 1913)
as being a troublesome bloodsucker at Tiberias.’”” There would appear to be some
confusion here, since the genus Tyichotanypus does not belong to the CERATOPO-
GONINAE, and the species included in it are structurally incapable of sucking blood.

Genus Foreipomyia, (Mg.) Kieffer.*
Forcipomyia (?) bipunctata, L. var.

One ¢g, Mount of Olives, 1.vii.1918, in Kaiserin Auguste-Viktoria Stiftung, on
window.

* No species of this genus is actually known to suck blood.

116 MAJOR E. E. AUSTEN.

The only obvious differences from British examples of F. bipunctata, L., exhibited
by the above specimen are that the hair on the scutellum is paler and perhaps longer,
that clothing the distal extremity of the abdomen paler, and that covering the wings
apparently shorter, while the knobs of the halteres are dead white without a tinge
of yellow.

Family CULICIDAE.

Of the mosquitos collected and bred by the author during 1917-18, a considerable
number were subsequently destroyed by Psocids. The material actually brought
home includes representatives of 20 species, all but two of which, however, have
already been recorded from various localities in Palestine by Captain P. J. Barraud,
in a paper published by him in this Bulletin a few months ago.* It will therefore
suffice to mention the additionst to Captain Barraud’s list, which both belong to
the genus Culex, and are as follows.

Culex tritaeniorhynchus, Giles.

One Q, bred, 9.viii.1918, from larva in floating débris: R. Auja, Khirbet Hadrah,
6 miles N.-E. of Jaffa.

Culex medestus, Fic.

Two 99, bred, 14,20.v.1918, from larvae in marsh at Tel Abu Zeitun, near
Jerisheh, 5 miles N.-E. of Jaffa.

It may be added that bionomical notes on various species of Anopheles will be
found in the author’s paper entitled ‘‘ Anti-Mosquito Measures in Palestine during the
Campaigns of 1917-1918,’{ although it now appears that in certain cases the
nomenclature there used is in need of revision. Thus—

A maculipennis, Mg., should be A. maculipennis, Mg. var.
A. sinensis, Wied., should be A. hyrcanus, Pall.

A. palestinensis, Theob., should be A. superpictus, Grassi.
A. turkhudi, Liston, should be A. multicolor, Camb.

Family SIMULIIDAE.

Genus Simulium, Latr.
Simulium flavipes, sp. n.

g.—Length (4 specimens), 2 mm.

Black ; anterior border of dorsum of thorax with a pale neutral grey patch on each
side, a similar patch on hind border in front of scutellum ; antennae vinaceous cinnamon
or light pinkish cinnamon ; genitalia indistinguishable from those of S. angustitarsis,
Lundstr.§; legs Naples yellow or pale buff yellow, tarsi, tips of tibiae and of hind
femora mummy-brown or sepia-coloured, front tarsi slender, not at all expanded, first
joint of hind tarsi strongly incrassate.

Head: palpt mummy-brown. Thorax: front border of dorsum clothed with
minute, appressed, glistening yellowish hairs; meso- and sternopleurae neutral
grey or light neutral grey. Abdomen: hair on abdominal scale pale yellowish.
Halteres cream-coloured. Legs: front and middle coxae (at least in some specimens)

* “ Mosquitos Collected in Palestine and Adjacent Territories,’ by Captain P. J. Barraud,
BE.Z.S., F.E.S.: Bull. Ent. Res., xi, pt. 4, pp. 387-395 (March 1921).

j For the identifications of these the author has to thank Mr. F. W. Edwards.

¢ Trans. Soc. Trop. Med. and Hygiene, xiii, no. 4, pp. 47-60 (November 1919).

§ Figured by Edwards, Bull. Ent. Res., vi, pt. 1, p. 24, fig. 1, 7 (June 1915), as those of
““S. aureum.”’

THE BLOOD-SUCKING DIPTERA OF PALESTINE. W7

with a dark brown or blackish spot or streak on posterior surface ; middle femora
with extreme tips light mummy-brown on upper surface ; hind legs clothed with
fine yellowish hair (tips of tarsi with brownish hair) ; hind tibiae strongly expanded
towards distal extremities ; last four joints of front tarsi together approximately
equal in length to, or slightly longer than first joint ; first joint of hind tarsi somewhat
lighter towards base, second joint short but without any noticeable dorsal excision,
combined length of last four joints of hind tarsi equal to slightly more than half
length of first joint.

Wadi el Kelt, Jordan Valley, near Jericho: type and three para-types, 1.vi.1918,
forming part of a number of ¢3 of same species, dancing in small swarms in dry
portion of Wadi bed, 5.30-6.30 p.m. At the same time 9° of Simulium equinum, L.
(see below), were abundant in the ears of horses picketed close by, on top of the
Wadi bank.

The species just described is allied to the Algerian Simulium beckeri, Roubaud
(Bull. Mus. d’Hist. Nat., xii, p. 520, 1906), but is apparently distinguishable, inter
alia, by its somewhat larger size, and by the leg markings, e.g., by the presence of
dark brown or brownish tips to the front and middle tibiae, and by the proximal
extremity of the hind tibiae being entirely yellowish (7.e., without a blackish ring
at the base). Szmulium flavipes is also closely allied to an undetermined Ethiopian
species (represented in the National Collection by five specimens from Zomba,
Nyasaland Protectorate), in which, however, the dark markings of the femora and
tibiae are more strongly developed.

Simulium equinum, L.*

Seven $4, Tel Abu Zeitun, near Jerisheh, 5 miles N.-E. of Jaffa, 2.v.1918—
part of small swarm dancing at foot of the Tel, near the marsh, 6.15 p.m.; 3 99,
Jericho Plain, 15.iv.1918, “‘ biting horses’ ears in morning and evening, all over
the Plain” (Capiain Kendle, Australian Veterinary Corps); 12 99, Jericho
Plain, 19.iv.1918, “‘in large numbers, attacking insides of horses’ ears st Toe
Hadrah Dam, R. Auja, 29.iv.1918, “flying low over the water, apparently ovipositing
on green slime close ta the water’s edge, 6.15 p.m.” ; 1 @ (pale-legged variety),
Jericho Plain, near Wadi el Kelt, 31.v.1918, “‘ in horse’s ear in evening.”

In the Wadi Hamis, near Ain Kanieh (about 11 miles N.N.-W. of Jerusalem),
on 8.111.1918, at a spot where the shallow water was flowing very swiftly, a greenish
Simulium larva was present in myriads on the stones in the bed of the stream. No
adults were seen, and under the circumstances it was impossible to attempt to breed
out any of the larvae. Close to the spot referred to, a number of Simuliwm larvae,
apparently belonging to three different species, were also found on the leaves of a
submerged piece of reed. Some of these latter larvae were collected and taken back
to quarters, and although the majority of those brought back died within two days,
two were observed on 10.iii.1918 to have pupated. One of the pupae (examined
and sketched as well as possible with the aid of an ordinary platyscopic lens) was found
to have on each side six slender respiratory filaments, apparently arising separately
from what looked like a long, narrow, whitish pad, the whole arrangement, except
as regards the thinness and length of the filaments, being similar to that of the pupal
respiratory organs of S. equinum, L., as recently figured by Edwards.

On 19.iv.1918 in the Wadi el Aujah (63 miles N. of Jericho), a rushing stream
of about five yards in width, Szmulium larvae were abundant on the stones. In
fact, just as in the Wadi Hamis six weeks earlier, so numerous were the larvae on
some of the stones that the latter looked as though they were hairy.

* For the definite determination of this species, the writer is indebted to Mr. F. W. Edwards.
{ F. W. Edwards, Bull. Ent. Res., xi, pt. 3, p. 236, fig. 6 (December 1920).

118 MAJOR E. E. AUSTEN.

At Hadrah Dam, R. Auja, on 29.iv.1918, when, as already recorded, a @ of
S. equinum was taken apparently ovipositing, the stones in the bed of one of the sluices.
were seen to have numerous small Simulium larvae on them.

No adults of Simulium were observed in Palestine in 1917, but on 9th June in
that year larvae were found on stones in running water in the Wadi Shanag (the
upper part of the Wadi Ghuzze), at Bir Esani, by Lt.-Col. Richardson, D.S.O.,
R.A.M.C. Similarly, in the Wadi Ghuzze itself near Tel el Fara, in a rapidly flowing
channel issuing from a pool, the writer noticed a few Simulium larvae on stones,
on Sist July 1917... Uherevean be little doubt that several species of Simulium, in
addition to the two recorded above, occur in Palestine, and that the genus will be
found to exist wherever the presence of running water affords the conditions necessary
for breeding.

Family PSYCHODIDAE.

Genus Phlebotomus, Rond.
Phiebotomus papatasii, Scop.

Probably abundant during the hotter months in every town and village in
Palestine, besides being widely distributed in the open country.

The small amount of material of this species which the author succeeded in bringing
home in safety consists of :—2 $3, 4 99, Deir el Belah, 8 miles S.-W. of Gaza,
13.viii.1917, in corners of an “‘ E.P.” tent, 10.30 a.m.; 1 2 (abdomen distended by
eggs), Deir el Belah, 22.viii.1917, in circular tent (alongside earth bank surmounted
by cactus hedge) in which a case of “ sand-fly (Phlebotomus) fever” had occurred ;
1 9, Deir el Belah, 8.ix.1917—both sexes numerous in upper corners of EP, tent,
used as office by 14th Coy. (A.T.) R.E., in palm grove at No. 14 Well; 2 3d, 2 2),
Bir ez Zeit (13 miles N. of Jerusalem), 22.vii.1918, in house—at least 5 2° seen in
dark, cellar-like basement of one house, and a number more specimens of both sexes
(one @ full of partly digested blood) in a dark, open cupboard in a room on first
floor of another dwelling ; 1 9, Et Tireh (about 5 miles N.N.-E. of Ludd), in corner
of dark granary; 1 9, Mulebbis, 2.viii.1918—with another specimen, in corner
of packing shed at orange grove.

More than one of the females enumerated above, which has the wing fringes,
especially those on the costal margins, decidedly dark, apparently belongs to
Newstead’s “ Dark form.’”*

In July and August 1917, the occurrence of a number of cases of sand-fly fever
in the E.E.F. at Deir el-Belah caused attention to be directed to Ph. papatasii, which
was at that time common in many places in dug-outs and tents, in the case of the
latter, especially such as were pitched in the palm groves near the wells or close
to banks of earth crowned with cactus hedges. Efforts to find breeding places
failed, though possibly lizard holes, more particularly the burrows of Agama stellio, 1
(“ Hardun” of the Arabs), the large repulsive-looking lizard so common about
cactus hedges in Palestine, may be among the sheltered retreats serving the insects
as nurseries. At Deir el Belah, 13.viii.1917, at 10.30 am., the writer found
Ph. papatasii abundant in an E.P. tent, situated in a garden close to a well and
occupied as sleeping quarters by enginemen working the pump at the latter.” The
insects were in numbers in the upper corners of the tent, sheltering behind the
valances and at the back of framed photographs hung across the corners ; there were
also several inside helmets and coats hanging on the,tent-pole. Both sexes were
present, 99, the majority of which were gorged with blood, perhaps slightly
predominating ; over 20 specimens were caught in tubes with little difficulty. At

* Newstead, Bull. Ent. Res., ii, p. 73 (1911).

THE BLOOD-SUCKING DIPTERA OF PALESTINE. 119

Jericho, in May—June 1918, Ph. papatasii was fairly common, and in the compound
of the Pilgrim’s Hospice at 4.45 a.m. on 7.vi.1918, the author was bitten on the
back of the hand by a @ of this species, in broad daylight. On 2.x.1918, when
occupying a tent on a spur of Mt. Carmel, above Haifa, the writer observed a 3
Ph. papatasvi inside his mosquito net.

Phlebotomus minutus, Rond., var. africanus, Newst.

The following are the data with regard to the few specimens of this insect brought
back by the author :—1 3, near Jerisheh, 5 miles N.-E. of Jaffa, 7.v.1918, in
author’s tent, 8.0 a.m.; 1 9, same locality and place, 15.v.1918, 7.15 p.m.; 2 $d,
2 99, near Yahudiyeh, about 8 miles E. of Jaffa, 24.viii1918, in E.P. tent in
olive grove; 1 g,6 92, and3 34, 7 29, near Jerisheh, 26.viii. and 3.ix.1918, behind
hanging valance of E.P. tent, in open country.

In the Jaffa district, at any rate during the period July-September 1918,
Ph. minutus var. africanus was generally to be found sheltering in numbers behind
the hanging flaps or valances in E.P. tents, while, if the settlement of Wilhelma
may be taken as a criterion, it was also common in villages. On 30.vil. 1918, the writer
was informed by Captain (afterwards Major) W. F. Corfield, D.A.D.M.S., 54th
Division, that the fly was abundant in all the houses in Wilhelma, and that in the
room used as the informant’s mess, 40 or 50 specimens were sometimes seen, or
were caught on the lamp, in one evening. On making a search in the office of the
A.D.M.S., 54th Division, in Wilhelma, the writer found two examples of what
appeared to be Ph. minutus var. africanus, while in the cellar-like basement of the
same house several more specimens of this Phlebotomus, including a female gorged
with blood, were seen and caught. Since every house in Wilhelma is provided
with a basement, the walls of which, being built of rough stone, are full of cracks
and crannies forming ideal breeding and sheltering places, the local abundance
of these ‘“‘ sand-flies ” was scarcely surprising, while, at any rate under active service
conditions, effective prophylactic measures were of course impossible.

When collecting specimens of Ph. minutus var. africanus in their favourite
retreats in E.P. tents, it was noticed that on first being uncovered they are often,
or usually, so completely motionless that they appear to be dead. On _ being
disturbed, however, they become very active, either jumping from side to side
after the manner of Ph. papatasii, or just as frequently taking wing at once. Placed
in tubes loosely plugged with cotton wool, these little flies frequently burrow into
the wool like minute mice, and then remain perfectly motionless. In the Jerisheh-
Wilhelma region, in the summer of 1918, occasional specimens of Ph. papatasi
were seen or caught in company with Ph. minutus var. africanus, though the latter
appeared to be far more numerous. Like Ph. papatasit, Ph. minutus var. africanus
sometimes shelters among clothing hanging on tent-poles; thus, near Jerisheh on
- 7.1x.1918, the author took 1 g and 3 9° of this variety among the folds of a Burberry

suspended from the pole in his tent. None of these females had blood in them,
and no specimens of Ph. minutus var. africanus were ever observed to bite; in fact
the finding in the basement of a house at Wilhelma, as mentioned above, of a single
female gorged with blood was the only evidence obtained that the present variety
is actually a blood-sucking fly. It only remains to add that whenever a series of
specimens was collected, females were found to be much more numerous than males,
and that a single male taken by the writer, 26.1x.1918, in his tent at Tul Keram,
appeared to belong to the variety under discussion.

Writing from observations made during a five weeks’ visit to southern Galilee
and Syria in October and November 1912, Dr. N. Annandale says* :—‘‘ By far the

* Journ. Asiatic Soc. Bengal, ix, pp. 44-45 (1913).

120 MAJOR E, E. AUSTEN.

most troublesome blood-sucking flies at Tiberias and Nazareth in October are the
so-called sand-flies of the genus Phlebotomus (fam. Psychodidae). They occur in
large numbers in every house, concealing themselves during the day in ceilings or
dark corners to which they retire shortly after sunrise, and commencing their
onslaught, which is continued until they retire, at sunset. Although I was unable
to find the larvae, I obtained indirect evidence that they breed in half-dried algae
just above the water level on the sides of open cisterns. Miss S. L. M. Summers,
of the London School of Tropical Medicine, who has been kind enough to examine
the adult specimens I collected, finds only two species (Ph. papatasi [sic], Scop., and
Ph. minutus, Rond.), among them, thus confirming the preliminary diagnosis made in
the field. Col. Alcock tells me that he found the same two species, and them only,
in a large collection from Aleppo. Phlebotomus apparently occurs at Tiberias
practically throughout the year, but at Damascus, in which it is troublesome in
summer, it had entirely disappeared before the end of October. I did not obtain
Ph. minutus at Nazareth, and at Tiberias it was much less common than Ph. papatast.”

Genus et sp. incert.

At Bir Esani, 24.x.1917, in the Wadi Immalaga near its mouth, on a ladleful
of water just taken from among bulrushes, the author found a small Psychodid
with a distinct proboscis and slightly spotted wings. Most unfortunately the
specimen, which was quite unlike anything seen before or since, was blown away
before it could be secured.

Family MUSCIDAE.
Genus Philaematomyia, Austen.

Philaematomyia crassirostris, Stein.
Musca crassirostris, Stein, Mitt. Zool. Mus. Berlin, ii, p. 99 (1903)

Philaematomvyia insignis, Austen, Ann. & Mag. Nat. Hist. (8) iii, p. 298, figs. iit
(1909).

Eleven ¢3, 11 99, Latron, close to Jaffa-Jerusalem road, 5.ix.1918; 1
Tul Keram, 26.ix.1918 (Lt.-Col, (temp. Col.) E&. P. Sewell; C.M.G, DS:
R.A.M.C.).

~~ Although from the moment of entering Palestine the writer kept a sharp look-
out for this widely-distributed species, it was not met with at all during 1917, and
even in 1918 it was not observed until 5th September, when the author took it in
abundance on his own person, at Latron. Had cattle been examined, the fly would
very possibly have been encountered earlier.* Patton and Cragg,+ writing of
Philaematomyia insignis, Austen (= Ph. (Musca) crassirostris, Stein), as observed
by them in Madras, state that flies of this species “ feed almost exclusively upon
Cathlesand?. % fe only occasionally bite human beings.’”’ The present writer’s
experience tends to support this statement. At Latron on the date mentioned he
took both sexes of Philaematomyia crassirostris in numbers on his puttees (24
specimens—l1 34, 13 92—were caught without difficulty in a killing tube), all the
insects being actively engaged in probing the material with their proboscises, evidently
in search of nutriment ; while, although the writer’s arms were bare and perspiring,
only a few specimens settled upon them, and only one of these latter flies (a 9)

2,
0.

*In Galilee, in October 1912, according to Dr. N. Annandale, the present species was “‘ by
far the commonest blood-sucking fly on horses and cattle’ :—-cf. E. Brunetti, ‘‘ Some Noxious
Diptera from Galilee ’’ (Journ. Proc. Asiatic Soc. Bengal, ix, p. 43, 1913).

+ Annals of Tropical Medicine and Parasitology, v, p. 518 (1912).

THE BLOOD-SUCKING DIPTERA OF PALESTINE. 12%

attempted to bite. It was felt at the time that so unmistakable a predilection for
settling upon and feeding from a hairy surface could scarcely be without significance.
The ° taken in the act of biting the author’s arm caused a sharp, pricking sensation,
which lasted for several minutes, although there was no visible blood at the bitten
spot. The 2 caught at Tul Keram by Colonel Sewell bit its captor on his bare leg
at 7.30 a.m., while he was dressing in his tent ; though disturbed before it had time
to draw blood, the fly left a mark upon the skin.

The greyness of the 99 of this species in life is very noticeable ; male specimens
do not appear nearly so grey. Another remarkable characteristic of this fly is the
softness of its body ; when pinning the specimens taken at Latron, which did not
appear to be newly-emerged, the author found it difficult to impale them on No. 20
pins, without crushing in the dorsum of the thorax, and the contrast in this respect
between Ph. crasstrostris and Musca domestica or M. autumnalis was most striking.

Genus Stomoxys, Geoff.
Stomoxys calcitrans, Linn.

One 3, 2 99, Deir el-Belah, 8 miles S.-W. of Gaza, 7,20.iv.1917 ; 1 9, Jaffa,
26.11.1918, in house ; 1 9, Wadi Hanein, near Richon le Sion, 9.iv.1918; 1 3 1 o
Deiran, 11.iv.1918, iv cozti.

Generally distributed. Brunetti (loc. cit.) states that in October 1912 this
species was met with in houses at Nazareth and Tiberias by Dr. N. Annandale,
who writes (7bid.) that it was ‘‘ also seen commonly on cattle.”

Genus Lyperosia, Rond.
Lyperosia irritans, Linn.

Fairly common in Wadi Ghuzze, near El Shellal, 11.xi.1917, when a number of
specimens settled on author’s sleeve, but did not attempt to bite; 1 ¢, 3 99,
Ain es-Sultan, near Jericho, 22.iv.1918, on horses.

[Lyperosia minuta, Bezzi.—Brunetti (Journ. Proc. Asiatic Soc. Bengal, ix, p. 48,
1913) records a single 9 of this species as having been taken at Tiberias, in October
1912, by Dr. N. Annandale, who writes (loc. cit.) :—‘‘ The specimen was caught
biting my hand at night. What I take to be this species is very troublesome,
especially in the early morning and at sunset, on the shores of the Lake of Tiberias,
easily piercing ordinary flannel with its proboscis. The wound is not very painful

oe al

and does not as a rule become inflamed.”’}

Family HIPPOBOSCIDAE.

Genus Hippobosca, Linn.
Hippobosca equina, Linn.

One 9, near Jericho, 6.iii.1918 (Captain (acting Lt.-Col.) W. J. Dale, O.B.E.,
R.A.V.C.); 1 9, Deir el-Belah, 8 miles S.-W. of Gaza, v.1917:; 1 QO, near
Jerisheh, 5 miles N.-E. of Jaffa, between 1.v. and 8.v.1918; 1 3, 1 9, Mulebbis,
21.v.1918, on cow (Captain C. Searle, M.C», R.A.M.C.); 1 3, Jericho Plain,
26.v.1918, on horse; 3 99, Mount of Olives, 26.v.1918, on horse (Captain (acting
Lt.-Col.) Dale); 1 3, 1 9, Mount of Olives, 20.vi.1918, inside Kaiserin Auguste-
Viktoria Stiftung; 1 g, Tul Keram, 27.ix.1918.

Brunetti (/oc. czt., p. 44) mentions that in October 1912 specimens of this species.
were taken by Dr. N. Annandale at Tiberias, Nazareth, and Kefr Kenna, “ sucking
blood of horses.”” The collector states (loc. cit.) that in Galilee H. equina is “ very
common on horses and cattle.”’

122 MAJOR E. E. AUSTEN.

Hippobosca capensis, v. Olf.

One 3, 1 9, Deir el-Belah, 8 miles S.-W. of Gaza, v.1917; 1 4 near Kefr
Insha, about 20 miles E. of Jaffa, 21.v.1918, settled in ear of author while he was
travelling in motor car; 1 3, Mount of Olives, 6.vii.1918, inside Kaiserin Auguste-
Viktoria Stiftung, on author’s neck; one specimen (sex uncertain, abdomen
missing), near Abud, 19 miles E. of Jaffa, 25.viii.1918, on author’s arm.

Hippobosca camelina, Leach.

One 3, near Jericho, 6.iii.1918 (Captain (acting Lt.-Col.) W. J. Dale, O.B.E.,
R.A.V.C.); 1 9, Tel es-Sultan, near Jericho, 21.1918, caught in horse lines,
with many specimens of Hippobosca equina, Linn. (Captain W. W. Averill, Auckland
Mid. Rifles); 1 3, 4 99, Deir el-Belah, 8 miles S.-W. ot ‘Gaza. 21x tol 7 som
camels.

It may be mentioned that a somewhat misshapen pupa (puparium), deposited
prematurely by one of the specimens last referred to, measures 7 mm. in length,
while the dimensions of a fully mature, empty puparium of H. camelina, obtained
at Biskra, Algeria, 3.ii1.1894 (Rev. A. E. Eaton) are—length, 6 mm., greatest
breadth, 5-4 mm.

Owing to the very large number of camels (some 30,000), chiefly from Egypt,
serving with the Egyptian Expeditionary Force in 1917-18, this species could have
been taken practically anywhere within the British lines in Palestine, at any rate
wherever there was a detachment of the Imperial Camel Corps or section of the Camel
Transport Corps.

Genus Lynchia, Weyenb.

Lynchia maura, Big.
Three $3, 3 99, Deir el-Belah, 8 miles S.-W. of Gaza, 23.vii.1917, on carrier-
pigeons belonging to Army Signal Service, R.E.

In the warmer parts of the earth, including southern Europe, this well-known
parasite of the domestic pigeon and disseminator of the haematozoon, Haemoproteus
columbae, Celli & Sanf., is nowadays to be found on its host practically wherever
the latter occurs. The Lynchia maura series already in the British Museum (Natural
History) includes specimens from the Canary Islands (Orotava, Teneriffe) ; N. Nigeria
(Hadeija) ; Nyasaland Protectorate (Kanyenda, Dwangwa R., W. Nyasa) ; Union
of South Africa (Cape Town, and Mowbray, Cape Province; Pretoria and Onder-
stepoort, Transvaal) ; Mauritius; Mesopotamia (Qurnah, R. Tigris) ; India (Deesa,
Ambala, Bangalore) ; Selangor (Kwala Lumpur) ; Jamaica (Spanish Town) ; and
Brazil (Para). Knab (Insec. Inscit. Mens., iv, 1916, p.-3), who gives records ranging
from Iowa to Southern Brazil (S. Paulo), states that the species “ is widely distributed
in America,” and that recently it.“ has made its appearance in the Hawaiian Islands.”

Apart from the dissemination of haematozoa, pathogenic or otherwise, by Lynchia
maura, these flies, several of which may occur on the same bird, are prejudicial to
carrier-pigeons by disturbing their rest. It was found in Palestine that an infested
pigeon on returning from a flight, instead of at once seeking its resting place, as these
birds usually do, would alight on the floor of the loft and proceed to stamp and
peck itself.

Genus Lipoptena, Nitzsch.

Lipoptena caprina, sp. n.

39.—Length, $ (7 specimens), 3-2 to 3-8 mm. (from anterior margin of clypeus
to posterior margin of scutellum, 2 to 2-2 mm.), 9 (3 specimens), 3-8 to 5mm. (from
anterior margin of clypeus to posterior margin of scutellum, 2:4 mm.) ; width of
head, ¢, 1 to just over 1 mm., 9, 1-25 mm.

THE BLOOD-SUCKING DIPTERA OF PALESTINE. 123

Dorsum of thorax (in dried specimens) shining mummy-brown ; chitin plates on
dorsum of abdomen small (first three plates in 2 minute) ; entire dorsum of abdomen
of 3 from posterior margin of basal segment backwards inclusive, except greater part
of the four plates of chitin and area immediately in front of last plate, thickly clothed
with relatively long, recumbent, cinnamon-rufous hair ; corresponding area of dorsum
of abdomen of 2 clothed for most part with very short hair.

Head: dorsal surface, including antennae, ochraceous-tawny, vertical triangle
(ocelligerous plate) dark brown or dark mummy-brown, nearly semi-circular and
extending much further forward than in L. cervi, L., frontal stryipe cinnamon-brown,
sepia-coloured or light mummy-brown, about half as broad again as long; each
inner orbit at its widest equal to or slightly exceeding extreme breadth of correspond-
ing eye; clypeus generally with a more or less distinct, isolated, pit-like depression
in middle line, midway between pit at posterior end of median longitudinal groove
and its hind margin, a dark brown horseshoe-shaped mark (more or less complete
or widely interrupted in middle line), usually fairly well defined, and with forwardly
directed concavity, encircling pit at end of median longitudinal groove, each arm
of the horseshoe running along inner edge of corresponding antennary pit, but not
reaching front margin of clypeus, a second, narrower, dark brown, curved mark,
interrupted in middle line by posterior pit-like depression, behind horseshoe and mid-
way between it and posterior margin of clypeus, arms of posterior curved mark not
extending so far forward as those of horseshoe, area adjacent to pit at posterior
end of median longitudinal groove brownish ; palpi dark brown, short; hair on
ventral surface of anterior border of head brownish at base, glistening ochraceous-
tawny towards distal extremity. Cephalic chaetotaxy: one bristle close to inner
upper angle of each orbit, on a level with posterior ocelli ; two bristles side by side
on each inner orbit,in a row extending obliquely forwards and inwards on a level with
upper margin of eye; one bristle (occasionally two bristles) on inner margin of each
orbit close to upper boundary of clypeus. Thorax: dorsum clothed with hair and
bristles of moderate length, dark brown at base, glistening cinnamon-rufous towards
their distal extremities ; middle line of mesonotum bordered on each side with a
curved row of bristles, commencing anteriorly a little in advance of hind margin of
humeral callus ; humeral calli each with six or seven bristles, postalar calli each with
three bristles ; lateral area of mesonotum on each side clothed fairly thickly with
bristles, of which those forming a transverse row on upper surface of protuberance
in front of base of each wing-stump are stouter and recurved ; scutellum sometimes
showing a pit-like depression (perhaps due to post mortem shrinkage) near each
lateral angle, sometimes also with a similar depression in middle line ; hind margin
of scutellum with six bristles. Each half of mesosternum roughly quadrate in outline
when seen from below, considerably larger than corresponding half of metasternum,
and closely beset with very short dark brown bristles, those on hind margin, apart
from usual long, hair-like bristle in front of socket of middle leg, larger and stouter
than elsewhere; short bristles on metasternum smaller and fewer than those on
mesosternum, though in this case also bristles on hind margin are stouter than
remainder. Abdomen: dorsum of g with basal segment of usual type, followed
in middle line by four small, transversely elongate plates of shining dark brown
chitin, widely separated by pinkish buff or cinnamon-buff integument ; transverse
diameter of last two plates about the same (0-6 mm.), but last plate considerably
deeper (7.¢., longer when measured from front to rear) than penultimate, the two
anterior plates very small, one-third or considerably less than half the size of the
penultimate, basal segment, except hind margin, clothed with short, appressed, dark
brown hair; venter cinnamon-buff, with a large horseshoe-shaped, dark neutral
grey mark not extending to distal extremity, and entire surface thickly clothed with
short hair, similar in colour and character to that on dorsum; dorsum of abdomen
of 2 with basal segment similar to that of 3, followed in middle line by four plates
of chitin widely separated by light ochraceous-buff integument, the terminal plate,

(3442) K

124 MAJOR E. E. AUSTEN.

consisting of dark brown chitin, situate at bottom of notch or depression in hind
margin of abdomen, and about equal in size to corresponding plate in 4, remaining
plates very small, light mummy-brown in colour and transversely elliptical or elliptical
oval in shape, the penultimate plate and the plate immediately following the basal
segment between one-third and one-fourth of the terminal plate in size, the ante-
penultimate plate considerably smaller than either of the two plates between which
it is situate; dorsum in 9 sparsely clothed with appressed, dark brown, chestnut-
brown or cinnamon-rufous hair, very short except on hind margin of basal segment
and on lateral margins of posterior half of abdomen, and, with exceptions stated,
much shorter than corresponding hair in ¢, each of the four median chitinous plates
with a more or less complete row of short hairs, varying in number, on or close to
its hind margin ; venter cinnamon-buff, fairly densely clothed with minute, appressed,
dark brown, chestnut-brown, or cinnamon-rufous hair. Legs, except tarsi, buff-yellow
or ochraceous-buff, front and middle femora brownish above towards distal extremi-
ties, anterior surfaces of front and middle tibiae also more or less brownish ; tarsi
cinnamon-brown or chestnut-brown ; bristles and hairs on legs dark brown to
cinnamon-rufous, stouter bristles dark brown at base, then paler.

Jerusalem and Ain Arik (10 miles N.N.-W. of Jerusalem): type of g, and 5 3
and 2 2 para-types, taken at Jerusalem, 29.vi.1918, on goats and kids; type O19;
and 1 ¢ para-type, caught at Ain Arik, 15.vii.1918, on a kid. In all cases the
insects were on the inside of the thighs; at Jerusalem on 29.vi.1918 several
specimens were found on one small kid. It may be noted that, in addition to
harbouring the parasite just described, the goats examined, which appeared to be
perfectly healthy, were also infested with ticks, specimens of which, apparently
belonging to two or three species, were numerous on the inside of the animals’ ears.

Among the microscopic preparations forming part of the National Collection
of Diptera is a Q Lipoptena, which is evidently conspecific with the specimens
enumerated above, and was taken on man at Snevce, Macedonia, in May 1918
.(Col. C. M. Wenyon, C.M.G., C.B.E., late ALMES2):

In general appearance Lipoptena caprina presents a close resemblance to L. ibicis,
Theob. (Second Report Wellcome Research Labs., Khartoum, 1906, p. 88, figs. 45-47),
which was described from specimens found on an ibex at Port Sudan, Anglo-Egyptian
Sudan. The new species agrees with L. ibicis with respect to size, etc., of the
abdominal plates of chitin and as regards the hairiness of the body, but is distinguish-
able by the row of bristles running obliquely across each inner orbit consisting of only
two instead of three or four, or sometimes even five bristles, and by the antero-
posterior diameter of the vertical triangle (ocelligerous plate) being if anything
slightly longer.

Owing to similarity of provenance, it might be reasonable to assume the identity
of Lipoptena ibicis, Theob., with L. chalcomelaena, Speiser (Zeitschr. f. syst. Hym. u.
Dipt., iv, p. 178, 1906), the typical series of which was obtained at Tor (Peninsula of
Sinai) on ‘‘ Capra caucasica’”’ (Capra aegagrus, Gmel. ?). While, however, the number
of bristles in the row running obliquely across the inner orbits as given by Speiser
for L. chalcomelaena agrees with what is found in L. ibicis, Speiser in describing
the abdomen of the male of his species gives no indication of special hairiness ; on
the other hand he describes the vertical triangle on the head as “ broad and short,”
whereas in L. ibicis, Theob., the antero-posterior diameter of the corresponding
plate is considerably longer than in L. cervi, Linn. On the other hand, in a
subsequent paper (of. cit., v, p. 354, 1905), Speiser, when giving additional localities
for his species, mentions that the majority of the specimens afterwards examined by
him were collected in Egypt (two on the shores of the Red Sea), and that all of these
bore a label with the MS. name of “L. hirta, Low.”’ This would suggest that if Speiser
has correctly identified these latter specimens as belonging to his own species,
Lipoptena ibicis, Theob., may still be a synonym of L. chalcomelaena, Speiser.

EXPLANATION OF “<PLATE IV.

. 1.—Wing of Culicoides tentorius, Austen, 9. x 73.
. 2—Wing of Culicoides odiiilis, Austen, 4. x 75.

. 3.—Wing of Culicoides newsteadi, Austen, 2. x 75.

BULL. ENT. (RESEARCH. Wor. XII. PART 2. PEATE IV.

SS ‘
rf

Fig. 1. Culicoides tentorius, Spwitys cme) foe

IN DOOX

E. FE. Austen ad nat. del.

+0
—~
*
~)
on

Fig. 3 Culicoides newsteadi, sp. n.,

* 3 -

nh —

a Ly eee oe 7 7

: Faience a
:

at pce i. > -
50 Fahl ereeia

125

COCCIDAE FROM THE SEYCHELLES.

By E. E. GREEN and F. Laine.

Pseudaonidia iota, sp. n.

Adult female pyriform, broadest across the meso- and metathorax ; cephalo-
thoracic area rounded in front, slightly contracted behind, where there is a well-defined
transverse grove ; abdomen tapering to the posterior extremity ; frons and margins
of thorax and abdomen with a few spiniform setae. Antennae rudimentary, each
bearing a single long straight seta. | Anterior spiracles with a small group of para-
stigmatic pores (fig. 1, A). Pygidium with a well defined, strongly chitinised,
pyriform, reticulated area, the lacunae oval or round, and disposed regularly ; the part
posterior to the anus, very heavily chitinised, obscuring the lacunae. Circumgenital
glands present, consisting of a continuous arch representing the united median and

Fig. 1. Pseudaonidia iota, sp. n.: A, adult, x 30; B, pygidiam, x 150; C, margin of pygidium,
< 250.

anterior lateral groups, with the posterior laterals separated by a short space; the
anterior laterals show signs of being in two rows. Dorsal pores numerous,
comparatively small, circular, arranged in longitudinal series on each side of the
pygidium, and extending on to the margins of the abdomen (fig. 1, B). Pygidial
margin with three pairs of lobes and numerous angular projections laterad, each lobe
notched on the outer side; two pairs of conspicuous claviform paraphyses, each
with a large separate circular knob like the dot of an 7 (fig. 1, C). Length, 1 mm. ;
greatest breadth about 0-75 mm.

Described from a single example.
On upper surface of leaf of Eugenia caryophyllata, Seychelles (P. R. Dupont).

The paraphyses of this species resemble those to be found in P. Jacinia, Brain
(Bull. Ent. Res., ix, 3, p. 207, March 1919), but the two species may be readily
separated by the absence of circumgenital glands in the latter.

Pseudaonidia aldabraca, sp. n.

Puparium of female more or less circular, brownish, partly overlaid with greyish-
white secretion. Diameter approximately 2 mm.; exuviae subcentral.

(3442) K 2

126 E. E. GREEN AND F. LAING.

Adult female broadly ovate, broadly rounded in front, bluntly pointed behind,
deeply incised on each side of the body immediately behind the cephalo-thoracic
area; broadest across the metathoracic area, 2.e., shortly behind the lateral incision.
The whole insect rigid, rather densely chitinous. Rudimentary antennae obsolete
or inconspicuous. Anterior spiracles with a small group of parastigmatic pores ;
posterior spiracles rather inconspicuous, with no pores. Segmental divisions of
abdomen marked by strong transverse folds which do not extend to the lateral
margins. A few longish spiniform setae at intervals along the margins of abdomen
(fig. 2, A). Pygidium not sharply defined ; its centre occupied by a large reticulated
area, the lacunae very irregular in size, form and disposition ; no circumgenital
glands; dorsal pores small, but numerous (fig. 2, B). Margin of pygidium with three
pairs of lobes ; the median pair larger and more prominent, bluntly conical ; second
and third pairs rather shallow, the apex of each nearer the inner side, more or less
conspicuously notched on the free edge. There is a suggestion of a fourth lobe in
the form of a sub-angular marginal prominence. Squames minute and inconspicuous }
a pair between the median lobes ; a second pair in the intervals between the median
and second lobes ; and one (possibly two) between the second and third lobes. There

Fig. 2. Pseudaonidia aldabraca, sp. n.: A, adult, x 30; B, pygidium, x 150; C, margin of
pygidium, x 225.

are three conspicuous crescentic incrassations, within the margin on each side, »
associated with short but rather stout paraphyses which extend inwards from the
interval between the median and second lobes and from the inner side of the third
and of the rudimentary fourth lobes. There is also a pair of short and more or less
confluent paraphyses between the median lobes (fig. 2, C). Length, 1-25 mm.;
greatest breadth, 1 mm.

Described from a single example.

On bark of “‘ Bois d’Amande,”’ Aldabra Island, Seychelles (P. R. Dupont).

This species is near to P. tesseratus, dEmm., but differs principally in its
smaller size, and in the absence of the strongly cristate lateral margins of the
pygidium.

Aonidia obtusa, sp. n.

Female puparium consisting almost entirely of the enlarged nymphal pellicle,
transversely oval, flat, or very slightly convex, a narrow marginal area ornamented
with sutures running irregularly from without inwards and intertwining; colour
varying from pale to dark brown, often thinly coated with white powdery secretion
over a wide marginal area, leaving only the centre bare (fig. 3, A). Pygidium recessed,

COCCIDAE FROM THE SEYCHELLES. 127

the extremity scarcely projecting, deeply incised on each side near the base ;_ margin
with a pair of narrow median lobes between which are two narrow squames, laterad
are two squames, a lobe, three squames, and then three lobes ; squames slightly
fimbriate. The basal areas (beyond the incisions) each with three prominent angular
projections. There are two pairs of broad (but obscure) semilunar pores, occupying
the intervals between the median, second and third lobes, and communicating with
conspicuous tubular ducts (fig. 3, B). Breadth, 1-1-16 mm.; length, 0-75 mm.

Fig. 3. Aonidia obtusa, sp.n: A, nymphal pellicle, x 33; B, pygidial margin of same, x 225;
C, adult, x 50; D, pygidium of adult, x 150.

Adult female broadly ovate, 0:8 to 0-95 mm. broad, by 0-75 mm. long;
abdominal segments compressed but visible. Antennae rudimentary, composed of
a short curved spine situated on a tubercle. Mouth-parts strongly developed ;
area surrounding mouth and anterior spiracles more densely chitinised than the
rest of the body (fig. 3, C). Pygidium obtuse, without a vestige of lobes or other
processes ; and with several conspicuous sub-marginal pores (fig. 3, D).

On Verschaffeltia splendida, Seychelles (P. R. Dupont).

Four names may be added to the list of CocciDAE already recorded from the
Seychelles.

1. Geroplastes rubens, Mask.

On fern ( Acvostichum sp.). Widely distributed in the Australasian region.

2. Chionaspis subcorticalis, Green.

On tomatoes, Astove Island, and on Sida sp., Assumption Island. Hitherto
recorded from Ceylon only.

oo)

. Pinnaspis buxi, Bouché.

On Pandanus sevchellarum, Felicité Island; also on Areca catechu. An almost
cosmopolitan species.

128 E. E. GREEN AND F. LAING.

4. Diaspis (Aulacaspis) flacourtiae, Rutherf.
On Flacourtia. Previously known from Ceylon only.

As Rutherford published his description (Bull. Ent. Res., v, 3, Dec. 1914,
p. 259) without any figures, we take this opportunity of supplying the omission
(see fig. 4, A, B).

Fig. 4. Diaspis flacourtiae, Rutherf.: A, adult, x 50; B, pygidium, x 150.

The species differs from pentagona—which it closely resembles—in the relatively
larger and more prominent median lobes, and in the obsolescent or very small
lateral lobes of the pygidium. The dentiform first lateral lobe, which is a prominent
feature in typical pentagona, is altogether lacking in flacourtiae. The pores on the
lateral margins of the body are far more numerous and conspicuous in flacourtiae.

Rutherford makes no mention of the pathological effect upon the host-plant
that is noticeable in Ceylon, where the insect causes a stimulation of growth in the
sub-lying woody tissue, causing irregular and conspicuous swellings on the infested
areas of the branches of Flacourtia. This effect is noticeable only upon the older
and mature branches. When the smaller and younger branches are attacked, there
is no such tendency to an abnormal growth.

| pik)

ON THREE. NEW /SPECIES OF INDIAN BRACONIDAE.
ByvG. livre, FES.

The following descriptions. are based on material submitted for determination
by Mr. T. Bainbrigge Fletcher, the Imperial Entomologist, Pusa, Bihar.

Family BRACONIDAE.

Genus Microplitis, Fdrst. (1862).
Microplitis similis, sp. n.

Black; palpi pale; legs testaceous, hind coxae at base and hind femora
darker, apical joint of all tarsi fuscous; sides of abdominal segments 1 and 2
testaceous. Wings hyaline, nervures fuscous, stigma unicolorous, testaceous or
fusco-testaceous. Antennae of male as long as body, of female one-third shorter,
fusco-testaceous, scape rather darker and flagellum darker towards apex. Head
and mesothorax granulate; scutellum smoother, dull; metathorax rugose, with
indications of a longitudinal medial carina. Abdomen smooth and shining, only
first segment feebly striolate ; shield of first segment twice as long as medial breadth,
slightly narrowed towards base and rounded at apex; second segment without
noticeable impressed lines. Spurs of hind tibiae barely one-third as long as meta-
tarsus. Terebra very short. Length, 23-3 mm., expanse, 5-6 mm.

Cocoons pale tan colour, similar to those of .W. spectabilis, Hal.
Type 9, in the British Museum ; cotypes in the Pusa Collection.
A parasite of Agvotis vpsilon, L., from the following localities :—

BIHAR & Orissa: Pusa, 1 J, 4 9, 16.iii.1914 (type material) ; Sabour, 2 9 (H. L.
Dutt). BENGAL: Mokamah, 1 4, xii.1911 (D. N. Pal),,and 2 g; 18.xii.1911. (C. S:
Misra).

Very near M. spectabilis, Hal., indeed at first I considered it to be a form of that
species, but the invariably unicolorous stigma and feebly longitudinally striolate
first segment of the abdomen would appear to warrant its separation. . spectabilis,
a common European species, has the stigma determinately pale at the inner angle,
the first abdominal segment minutely punctuate, and the hind and middle tarsi
fuscous.

Microplitis eusirus, sp. n.

Black ; palpi pale; legs rufo-testaceous (fore and middle coxae and middle
femora except at apex fuscous, hind legs entirely black or blackish excepting
trochanters and tarsi towards apex, which are often rufo-fuscous). Sides of first and
second abdominal segments lighter in colour. Wings infumated, with the usual
dark blotch under the stigma ; nervures fuscous ; stigma unicolorous, dark fuscous.
Antennae stout, rufo-fuscous ; scape rufous at base ; each joint of flagellum centrally
marked with an impressed band which gives the antennae the appearance of having
twice as many joints as is actually the case (this character is noticeable in a lesser
degree in some of the European species). Eyes pilose. Head and mesothorax
densely and minutely punctuate; sutures of the mesothorax deeply and clearly
impressed ; scutellum rugulose. Metathorax coarsely rugose, with a medial trans-
verse centrally angulated carina, after which the metathorax falls suddenly away
(fig. 1, a). Abdomen smooth and shining; shield of first segment elongate, three
times as long as medial breadth, with parallel sides, slightly widened at base and

130 Ga D.. bynes

truncate at apex (fig. 1, a), apical tubercle not prominent. Spurs of hind tibiae pale,
less than one-third as long as metatarsus. Terebra very short. Length, 3}-4 mm., -
expanse, 7-8 mm.

Type 2 in the British Museum ; cotype in the Pusa Collection.

Brnar & Orissa: Pusa, 1 g, 2 9 and fragments of 2 others reared from Achaea
janata, L., 20.ix.1912 (H. L. Duit).

Fig. 1 (a). Propodeon and first abdominal tergite of Microplitis eusirus, sp. n., Q ; in the pro-
podeon only the major reticulations are shown; the point of view is at right angles to the
median carina, and the dorsal surface (unshaded) is seen considerably foreshortened ; only the
outline of the plate on the Ist tergite is indicated.

(b). Rhogas (Heterogamus) percurrens, sp. n., Q; detail of wing.
(c). Heterogamus dispar, Curt., 3; detail of wing.
(d). Heterogamus dispar, Curt., ©; detail of wing.

Genus Rhogas, Nees (1818).

The following species is, formally at least, referable to Heterogamus, Wesm.
(1838), but the difference between the first and second abscissae of the radius is so
slight that the insect might with almost equal propriety be placed in Rhogas, Nees
(fig. 1, 6). The genotype of Heterogamus, Wesm. (H. dispar, Curt.) is a rare and
extremely distinct species, and quite possibly Heferogamus may be a valid genus ; but
if so, it is not yet properly understood and its real differentiae have not been fully
signalised. I have not had before me sufficient material of the genotype to satisfy
myself on this point, and for the present treat Heferogamus as a subgenus. Itcan |
deserve no higher rank than this if its only criterion be the ratio between the first
and second radial abscissae.

ON THREE NEW SPECIES OF INDIAN BRACONIDAE. 131

Rhogas (Heterogamus) percurrens, sp. n.

°.—General colour of antennae,* body and legs pale ferruginous. The ocellar
triangle, but not the ocelli themselves, blackish. Mesonotum with a broad longi-
tudinal faintly infuscated band inside each notaulus and two narrow short ones on
the mid lobe anteriorly. Wings with costa, including basal three-fifths of the stigma,
concolorous with the body, apical two-fifths of stigma infuscated. Nervures mainly
darkened, but the second abscissa of the cubitus very pale. Abdomen a little
darkened posteriorly from about the middle of the fourth tergite. Venter all pale ;
sheath of ovipositor blackish brown. Antenna with joints subequal, about one-third
longer than broad. Thorax with transverse sulcus at base of scutellum crenulate
with about eight large punctures or pits. Sides of scutellum with seven to eight
short, stout, parallel ridges. Metanotum, apart from the usual divisions, smooth.
Propodeon, in the type, with the median keel not completely percurrent, reaching
back only to about two-thirds; the surface on each side of the keel irregularly
rugulose ; spiracle moderate, oval, a little over twice its length from the anterior
edge.

Fig. 2. Basal portion of antenna of (a) Heterogamus dispar, Curt.; (b) Rhogas (Heterogamus)
percurrens, sp. Nn.

In the wings (fig. 1, 6) the second cubital cell is a little, but quite perceptibly,
narrowed distally, the second abscissa of the radius being slightly inclined to the
third abscissa of the cubitus ; first intercubital nervure and second abscissa of the
radius subequal, the latter again much longer than the second intercubital. The
radius and cubitus strongly divergent distally. The stigma distinctly broad.

Abdomen with the median keel percurrent to the posterior edge of the third
tergite, sharply defined throughout its course, though broader on tergite one.
Throughout its course the mid keel is flanked by numerous subparallel longitudinal
ridges or rugae, these being continuous from tergite to tergite and extending even
to the anterior or basal one-third of tergite 4. The first three tergites are rigid,
and the sutures (especially the first) deep and distinct. The second suture (between
two and three) might almost be described as crenulate, as the sulcus is cut into little
pits by the percurrent rugae. Beyond the basal one-third of the fourth tergite
the surface is smooth ; only the smooth edge of the fifth tergite is visible.

Length just over 5 mm.; alar expanse, 9 mm.
Type a 9 in the British Museum.

* In the single antenna preserved, which is complete up to the 23rd joint (21st of the funicle)
the colour is uniformly pale ferruginous. without any suggestion of banding.

132 (EL at NALD

Brnar & OrIssA: Pusa, a unique 92 reared from Achaea janata, | pia fee) he 2 ys
(OC. Ke Dutt):

R. percurrens sp. n., has a strong but probably superficial resemblance to
Heterogamus dispar. So far as colour is concerned the resemblance is to the ¢ of
that species. In colour, however, dispar is notoriously variable. It is possible that
the darkening of the thorax and abdomen in percurrens owes something to the drying
up and discoloration of the underlying musculature of these regions.

H. dispar is easily separated from the present insect (a) by the basal antennal
joints, which are at least twice as long as broad (fig. 2) ; (b) in the wings (fig. 1, ¢, @),
the radial cell is longer and more acute, the second cubital cell “higher” (2, @),
the second abscissa of the radius being just shorter (2) or much shorter (3) than the
first intercubital. The same abscissa is again equal to (2) or much shorter than (3)
the second intercubital. The radius and cubitus are less abruptly divergent distally
and the stigma is narrower. In the abdomen the median keel is percurrent to the
posterior margin of the second tergite only in the material available for examination.

133

thi PTINEDY BEELILE EPRIGONOGENTUS :-GLOBULUM, SOLIER:
BREEDING IN ARGOL.

Bye nwuen.scorr, MAL Sep FES,

Curator in Entomology, University of Cambridge.

In March 1920 my attention was called by Dr. F. W. Dootson, University
Lecturer in Chemistry, to the fact that a beetle was breeding in numbers in a jar
of argol in the Chemical Laboratory of Cambridge University. The insect proved
on examination to be the introduced Ptinid, Tvigonogenius globulum, Solier,* a
form related to the household insect, Niptus hololeucus, to which it bears at first
sight a slight resemblance.

Argol is the crust or deposit which separates out in barrels of new wine. It
contains a high percentage of cream of tartar (potassium bitartrate), and most of
the pure stock of that chemical is prepared from it. The argol in which the beetles
were living was found to contain about 80 per cent. of potassium bitartrate. The
argol was a purplish-red powder of close consistency, and it came to the Chemical
Laboratory from London in a bag in 1913. It was placed in an earthenware jar,
tightly corked with a wide cork bung. The powder did not quite fill the jar, but a
very small air-space was left at the top.t Iam assured that the jar was not opened
from the time the powder was put in till early in 1920, when the argol was found to
be full of adults and larvae of the Trigonogenius. The cork, which I have twice
examined, has not been bored through or damaged by insects in any way, and it
fits so closely that it is almost impossible that the beetles can have got into the jar
down the side of the cork; nor have they been noticed anywhere else in the
laboratory. Probably, therefore, some of the insects were in the argol when it came
to the laboratory and have continued breeding in it all these vears. No other kinds
of insects were found in the argol. I am indebted to Dr. Dootson for most of the
foregoing particulars and for samples of the infested chemical.

That the beetles were nourishing themselves, not exclusively on the 80 per cent.
of potassium bitartrate, but at any rate partly on some of the ingredients forming
the other 20 per cent. of the argol, seems to be indicated by the following experiment,
carried out at Dr. Dootson’s suggestion. Some of the insects were placed on the
surface of about two inches depth of pure cream of tartar in a wide-mouthed corked
bottle, with plenty of air-space between the surface of the chemical and the cork ;
32 adults and 15 larvae were placed in this on 12th March 1920. Three and a half
hours later almost all the larvae had burrowed down into the white powder, but the
adults were still on the surface and showing signs of discomfort. Next day all the
larvae but one were below the surface, and also about 23 of the adults ; the remaining
adults were still on the surface and one was dead. No further observation was
made for nearly three weeks, when (on Ist April 1920) about 14 adults and one larva
were seen to be on the surface, while the other 18 adults and 14 larvae had all burrowed
some way down into the cream of tartar, and several burrows were visible against
the glass sides of the bottle. The bottle was not examined again for nearly a year,
when (on 15th March 1921) its contents were turned out, and all the insects were

* Described by Solier in C. Gay’s “‘ Historia de Chile,” iv, 1849, p. 464. I have retained Solier’s
original spelling of the specific name, though in some later works globulum is altered to globulus,
which is probably more correct. Solier called the species globulum, but at the same time named
a variety of it globosus. Why he gave the former name the neuter ending is difficult to say.
He may have intended globulum to be a noun, but in several dictionaries of classical and late Latin
which I have seen, the only form of the word is a late Latin noun, globu/us.

¢ In some of the argol which was put into a glass-stoppered bottle, filling it up to the stopper
and leaving no air-space, all the insects died.

134 HUGH SCOTT.

found to be dead ; there were four adults on the surface and 34 adults below, and
remains of ‘about cight larvae were found. It will be noticed that the adults were
six more in number than those placed in the bottle the year before, proving that
some of the larvae had succeeded in reaching the adult state ; most of them when
put in were fairly big. But the attempt to start a culture breeding in pure cream
of tartar failed. It should be mentioned that no moisture was supplied, but neither
was it, of course, in the original jar of argol, which was however far larger and
contained a very much greater bulk of chemical than the bottle used in this
experiment.

In the original jar of argol the insects were still present in large numbers in
April 1921. Series of adults and larvae have been preserved, but I have found
only one pupa, though samples of the argol have been looked through on several
occasions. This single pupa was found at about the end of March 1921. Mr. Michael
G. L. Perkins, who placed some of the argol in a vessel and kept the beetles present
in it under observation, obtained about 30 pupae in July 1920. He has also started
cultures of the insect in certain food-stuffs such as oatmeal and raisins, and, so far
as his observations have gone, he considers that the generations succeed one another
more rapidly in these food-stuffs than in the argol, in which substance the insects
appear to be reproducing themselves only at the rate of about one generation a
year. He hopes to publish the results of his investigations in the future.

The larva pupates in a cocoon formed of a feltwork of fine threads, secreted
by itself. The cocoon is not lined, and the feltwork is easily pulled apart with
needles into a loose tangle of threads. The feltwork appears whitish when the argol
powder is shaken away from it. Under a high power (2-inch objective) the threads
are quite colourless and transparent. Many full-fed larvae, pupae, and adults have
been found in these cocoons by Mr. Perkins and myself. Mr. Perkins sifted the
argol, to remove all foreign material from it, before he placed his observation culture
of the insects in it, thereby proving that the threads are actually produced by the
larvae. He thinks that other feltwork is produced in the burrows, besides that
actually used in the construction of the cocoons.

Trigonogentus globulum was originally described from Chile, but is very widely
distributed. In M. Pic’s Catalogue of Ptinidae (1912, Col. Cat., part 41, p. 9), it
is recorded from various parts of North and South America, Tasmania, and England.
Most of the known species of the genus are, according to the Catalogue, known from
Central or South America, and several of them are not known from elsewhere, so
that region may be their native home. Fowler & Donisthorpe (Col. Brit. Isl., vi,
1913, p. 147) state that it has occurred in corn mills and granaries in various industrial
centres in England. Champion (Ent. Mo. Mag., 1918, p. 40) records it as having been
found under timber and among wood-shavings in London, in company with Pétimus
tectus and Niptus hololeucus: and Potter (op. cit. 1919, p. 88) records it from old
cotton mills near Manchester, where it was attracted to baits of sugar It is not
mentioned in Reitter’s “‘ Fauna Germanica: Die Kafer ’’ nor in the addenda at the
end of the last volume of that work (Vol. V, 1916), so it had evidently not occurred
in Germany up till 1916 within the knowledge of the writers of that book. I have
not searched the literature further. Péinus tectus and Niptus hololeucus, the members
of the same family mentioned above, are, like Trigonogenius globuluwm, found in
various stored products. Ptinus tectus further resembles it in being almost cosmo-
politan in range. It may be recalled that larvae and adults of Niptus hololeucus
have recently been found in cocoa powder from a south German chocolate factory
(Rev. Appl. Ent., A.ix, 1921, p. 66).

135

® REVISION OF THE GENUS LOCUSTA, L. (= PAGHYTYLUS, FIEB,),
Witt A NEW THEORY AS TO THE PERIODICITY AND MIGRATIONS
OF LOCUSTS;

By. 5B» P." Uvarov, B.E-S.;

Assistant Entomologist, Imperial Bureau of Entomology.

CONTENTS.
PAGE.
I. Introductory.. nfs Re aN a ¥; x» ae J *eihoo
II. On the generic name Locusta, L. od ae ss oe sa loo
III. Locusta migratoria, L., and its forms .. 7 hg: ate oe ABT
IV. Locustana pardalina, Walk., and its phases .. ae x bdeh Oo
V. Systematic part <a 2% ae ~ 159

I. INTRODUCTORY.

The genus Locusta, L. (= Pachytylus, Fieb.) includes two of the most destructive
swarming locusts of the Old World: the widely distributed L. migratoria, L. (with
L. danica, L., and L. migratorioides, Rch. & Frm., as its forms; see below), and the
South African L. pardalina, Walk. The literature on the economics, biclogy and
especially on the means of control of these locusts is enormously extensive, but at
the same time their systematic arrangement is in considerable confusion, and
extremely contradictory opinions as to the mutual relationship of the different
so-called species exist among specialists. As a direct consequence of this, the field
research and control work of economic entomologists is apt to suffer through the
difficulty in getting a particular species properly named,* and thus it is often
impossible to make a comparison of the records as to the biology and control of
the same species in different countries.

Having had the opportunity of conducting, during the years 1911-14, extensive
field research work and control work on L. migratoria in the northern Caucasus
(Russia), and being a systematist, I could not fail to see at once that only very little
progress could be made without a definite solution of the question of the inter-
relation between L. migratoria and L. danica, which latter has been accepted
by many authors as a species distinct from migratoria, and as conspecific with it,
by others. The same question arose before the Turkestan Entomological Station
(in Tashkent) as soon as its staff began to work on L. migratoria.

Apart from my field work, which involved the study of immense series of living
specimens in all stages, I endeavoured to gather all reliable information as to the
distribution and local, individual and annual variability of L. migratoria and danica
in different parts of their range, and owing to the support of entomologists and
institutions throughout Russia and elsewhere, I managed to concentrate in my
hands extremely rich materials from the following sources : Turkestan Entomological
Station (V. j. Plotnikov); Astrakhan Entomological Station (N. L. Sakharov) ;
Natural History Museum in Kherson (J. K. Pachossky) ; Zoological Museum of the
Moscow University (Prof. J. Kozhevnikov) ; Caucasian Museum in Tiflis ; Zoological
Museum in Berlin. The following persons also contributed very valuable materials
and information: Prof. J. Shtchelkanovzev, E. Jatzentkovsky, V. Artsimovicz

* See, for instance, the interesting paper on the biology and control of the Malayan locust
by H. C. Pratt (Bull. No. 27, Dept. Agric. Fed. Malay States, 1915), who states that “ many
attempts to identify this Malayan locust have been made and correspondence has been entered
into with authorities throughout the world, but without success.’? He has been compelled to
call the insect, simply, Pachytylus sp.

136 B. P. UVAROV.

N. Ikonnikov, V. Boldyrev, L. Moritz, H. C. Pratt (Government Entomologist,
Federated Malay States), and many others. Thanks to this generous help, I was
able to come to certain conclusions on the question of migratoria-danica already
in 1915, but the War and other circumstances prevented me from publishing a paper
on it. When I arrived in London in 1920 and studied the exotic representatives
of the genus Locusta, I felt the necessity of revising my previous work and extending
its limits so as to include in it all known species and forms of the genus. This plan
proved to be a very productive one, since a far more definite idea as to the inter-
relations of different ‘‘ species’? has been thus reached. A study of the South
African L. pardalina, Walk., has been also accomplished in the British Museum, but
I could not do much with museum material only, and the most effective help in this
respect has been rendered me by the Division of Entomology, Pretoria, and especially
by Mr. J. C. Faure, who has sent to the Imperial Bureau of Entomology at my
request extensive series of specimens, together with most valuable information.

With regard to the biological observations here recorded, it is only thanks to the
help rendered me by my assistants, G.Vinokurov, Th. Gliniuk, the late G. Pirkovsky,
and others, that I could collect the necessary facts. My most sincere thanks are
due also to all the above-mentioned persons and the heads of institutions who have
lent me material and supplied information.

Il. ON THE GENERIC NAME Locusta, L.

The Linnean genus, Gryllus Locusta, includes 20 different species of locusts and
grasshoppers, belonging to about as many modern genera. There is no wonder,
therefore, that much controversy arose around the question as to the species to which
the Linnean name Locusta must be restricted now. This question becomes still
more complicated owing to the fact that Geoffroy (Hist. Ins. i, p. 396, 1762) applied
the name Locusta quite erroneously to the long-horned katydids (now called
TETTIGONIDAE, or, wrongly, PHASGONURIDAE), and has been followed in this mistake
by all continental European authors, till quite recently. There is, however, no
doubt, that Linné, who adopted the name Locusta from the old Roman writers who
applied it to swarming locusts, intended it to include those insects and the short-
horned grasshoppers generally. This view was accepted long ago by British authors,
and W. E. Leach (Edinburgh Encyclopaedia, ix, pt. 1, p. 120, 1815), though using
Locusta, Geoffr., for katydids, used at the same time Gryllus Locusta, L., for migratoria,
L., the latter being the only species described by him under this genus, and, therefore,
he actually has fixed it as genotype of Locusta, L. A few years later on, Samouelle
(Entomologist’s Useful Compendium, p. 218, 1819) followed Leach in restricting
the genus Locusta to migratoria, but he calls it wrongly Locusta, Leach, not Linné.
Stephens in 1829 (Cat. Brit. Ins.,i, p. 301, No. 2, sp. 3315) merely repeats Samouelle’s
interpretation of the genus. Even if we do not accept the genotype of Locusta, I
cited in 1815 by Leach, we shall find a most formal fixation of it in the British Ento-
mology of Curtis (iii, pl. 608, August 1836), who in describing Locusta christi, Curtis
(=danica, L..) said positively: ‘“‘ Type of the genus, Gryllus migratorius, L.” All
subsequent works on the same subject are, thus, of no importance, and the generic
name Pachytylus, Fieber, proposed in 1853 (Lotos, iii, p. 121) for migratora and
danica is a pure synonym of Locusta, L. The proposal of Rehn (Canadian Ento-
mologist, xxxili, 1901) to restrict the genus Locusta, L., to apricarius, viridulus and
biguttulus, which are included now in the genera Stauroderus and Omocestus, as well
as that of H. Krauss (Zvol. Anz., xxv, 1902, p. 539), who regarded tatarica, L., as
the genotype of Locusta, cannot be accepted in view of Ctrtis’ work, which was
overlooked by both these authors.

A REVISION OF THE GENUS LOCUSTA, L. oy

It is, therefore, in full accordance with what Linné meant by his genus Gryllus
Locusta, as well as with the formal laws of nomenclature, that migratoria, L., must be
regarded as the genotype of Locusta, L.*

+”?

Not less than 16 “ species ”’ have been described by different authors as belonging
to the genus Locusta, L. (=Pachytylus, Fieb.). This number, however, has been
reduced already by earlier revisers, who synonymised many species; but W. F.
Kirby in his Catalogue (Syn. Cat. Orth., iii, 1910, pp. 221-231) still mentioned
seven distinct species. My investigations, however, have clearly demonstrated the
variability of the species of Locusta to an extent far greater than might have been
anticipated, and my conclusion, which will be fully proved presently, is that only
two species can be distinguished, namely, migratoria, L., and pardalina, Walk.,
but the latter differs from migratoria in so many important characters that a new
genus is described below (p. 162) to include it, which I propose to call Locustana, g.n.

III. Locusta MicRATORIA, L., AND ITS FORMS.

Morphological Characters and Variability of migratoria and danica.

These two forms, if typical examples are studied, seem to be quite distinct from
each other in many morphological characters, and may be regarded, as has been done
by most authors, as two independent species. On the other hand, every extensive
collection includes specimens of Locusta that cannot be identified with certainty
with either migratoria or danica, but seem to represent intermediate forms. This
fact induced many authors to regard migratoria and danica as but extreme individual
aberrations of the same species.

In studving this question I tried, first of all, to analyse carefully and impartiallyt
all the external morphological characters of both forms, as given by different authors,
studying them on as extensive a series of specimens as possible.

After excluding all characters that at once proved to be too indefinite or simply
incidental, the following summary of differences between typical danica and migratoria
has been obtained :—

L. migratoria (fig. 1, C, D, E.)

Vertex convex, with a median lJongi-
tudinal keel; fastigium separated from
frontal ridge by an angular transverse
keel.

Pronotum relatively shorter and
broader in metazona, with a distinct
construction before the middle; fore
margin rounded ; hind angle rounded ;
median keel low, in profile straight or
even concave.

Elytrat relatively longer.
Hind femora relatively shorter.

L. danica (fig. 1, A; B.)

Vertex flat, without median keel ; no
transverse keel separating fastigium
from frontal ridge.

Pronotum relatively longer and more
compressed laterally, without or with
but feeble constriction before the middle;
fore margin angulately prominent ; hind
angle acute; median keel high, tecti-
form, convex in profile.

Elyira relatively shorter.

Hind femora relatively longer.

* Tam much indebted to Mr. J. H. Durrant for the help he has generously given me in the

solution of this question.

+ I must candidly confess that when starting my work I had only a very modest intention—
to find out characters for separating migratoria and danica, which I assumed beforehand to be
distinct specifically. The facts quickly destroyed my preconceived opinion and compelled me

to work deductively.

{ This important and rather striking difference between migratoria and danica was first noted
by the artist of the Zoological Museum in Petrograd, Miss O. M. Somina, who made drawings
of both insects for Mr. I. Shevyrev, and the latter drew my attention to it.

(3442)

Yr
ia

138 B. P.. UVAROV.

A study of long series of both forms showed that the differences in the vertex,
as given above, are quite unreliable, since the median keel of the fastigium proved
to be absent in one per cent. of the specimens of migratoria, as well as in about 7 per
cent. of danica; the transverse keel was found in 80 per cent. of migratoria and in
about 47 per cent. of danica; as for the flatness or convexity of the vertex, it is
too indefinite a character to be reliable. The only trustworthy characters are,
therefore, the shape of the pronotum and the relative length of the elytra and hind
femora. In order to avoid, as far as possible, any subjective judgment, I have
expressed these characters in terms of actual measurements. This is quite simple
in the case of the elytra and femora, since there exists an inverse proportion between
their lengths, and we may simply express the length of the femora as a percentage

Fig. 1. Locusta migratoria, L.: A, B, phase danica, L., 9 Turkestan ; C, D, ph. migratoria 3,
progeny of preceding specimen; E, ph. migratoria, typical ©, Astrakhan ; F, elytron.
Elytron natural size, remainder x 3.

of the length of the elytra. As for the shape of the pronotum, numerous measure-
ments have shown that the width of the metazona (measured between the shoulders,
i.e., at its widest), if expressed as a percentage of the length of the whole pronotum
(along the median keel), gives the best and most reliable impression of the actual
shape. The relative height and form of the median keel, which seem to be excluded
from consideration by using this proportion, is but a secondary character which
depends entirely on the relative length and width of the pronotum. If we imagine
that the short and broad pronotum of migratoria undergoes a lateral compression,
we may expect it to become longer, and its median keel higher and convex, especially
in the metazona, which should be most influenced by lateral compression ; there is
no doubt, therefore, that the shape of the median keel must be, and actually is, subject
to changes correlated with those of the length and width of the whole pronotum.

A REVIS{ON OF THE GENUS LOCUSTA, L. 139

Though it would be interesting and useful to give individual dimensions of all
the specimens measured, I refrain from doing so in order not to encumber this paper
with many pages of figures. I will give, therefore, only a general table of the chief
results obtained, which will be enough for our conclusions (see Table I).

TABLE I.

Showing Dimensions of different Phases of L. migratoria, L.

ee are Pronotal Proportion.* Femoral Proportion.t
g g | an KS) S a ieee
a) SS), Bol et) 8 Peel eel 8 ie Bee | us
= BD S = % a eh | cece ae é 1 es fe
i= = = 5 = ye] [ete < = = a> <
1. All migratoria .. AC mare lee 171 — — 0-89 | 0-71 | 0-18 | 0-80 | 0-53 | 0-40 | 0-13 | 0-46
2. All danica =e Sra mG 165 — 166 — 0-85 | 0:59 | 0-26 | 0-72 | 0-63 | 0-43 | 0-26 | 0-53
3. Palaearctic danica ate wa | LOS — 103 — 0-84 ; 0-59 | 0-25 | 0-72 | 0-63 | 0-43 | 0-20 | 0-52
4. Extra-Palaearctic danica aie 63 — 63 — 0-85 | 0-66 | 0-19 | 0-74 | 0-60 | 0-46 | 0-14 | 0-55
5. Stavropol province, 1912 se 25 25 — a 0-87 ; 0-71 | 0-16 | 0-79 | 0-53 | 0-42 | 0-11 | 0-45
6. do. do. 1913 ais 32 19 8 & 0-87 | 0-66 | 0-21 | 0:77 | 0-53 | 0-41 | 0-12 | 0-47
7. Valley of R. Kuma, 1913 Be 28 23 4 1 6-87 | 0-64 | 0-23 | 0-78 | 0-53 | 0-41 | 0-12 | 0-46
8. do. do. 1914 ae 36 95 8 3 0-87 | 0-65 | 0-22 | 0°76 | 0-54 | 0-42 | 0-12 | 0-48
9. Lac Vshivoye, 1912 .. zs iff) 10 1 — 0-80 | 0-74 | 0-06 | 0-78 | 0:48 | 0-41 | 0-07 | 0-45
10. do. dois A913) 2. ae 8 Z 4 2, 0-80 | 0-69 | 0-11 | 0-75 | 0-51 | 0-48 | 0-03 | 0-49
11. Konstantinovka, Stavropol prov.
191 |. Ne 3% — 10 — 6 4 0-77 | 0-68 | 0-09 | 0-72 | 0-55 | 0-45 | 0-10 | 0-50
12. Kalaus, Stavropol prov., 1913. . 3) 7 —- 2, 0-89 | 0-73 | 0-16 | 0-79 | 0-49 | 0-45 | 0-04 | 0-47
13. Valley of R. Terek, 1913 te 26 22 — 4 0-87 | 0-76 | 0-11 | 0-80 | 0-50 | 0-42 | 0-08 | 0-46
14, Valley of R. Syr-Darya, 1912.. 16 16 — —- 0-89 | 0-74 | 0-15 | 0-80 | 0-50 | 0-40 | 0-10 | 0-45
15. Mangishlak, Caspian Sea, 1911 10 10 — — 0-87 | 0-73 | 0-14 | 0-80 | 0-46 | 0-43 | 0-03 | 0-45
16. Korea, 1900 .. sus oe 8 — 8 — | 0-78 | 0-68 | 0-10 | 0-73 | 0-60 | 0-52 | 0-08 | 0-55
17. Smyrna .. are ae “A 14 — 14 — | 0°84 | 0-67 | 0-17 | 0:75 | 0-63 | 0-51 | 0-12 | 0-55
18. Azores .. ate “5 vs 20 — 20 — | 0-85 | 0-69 | 0-16 | 0-75 | 0:60 | 0-52 | 0-12 | 0-57
19. Congo Ae Ne 9 | — 9 — | 0-82 | 0-68] 0-14 | 0-75 | 0-59 | 0-50 | 0-09 | 0-54
migratorioides , |
‘20. S.W. Nyasa 4 -- os — | 0-94 | 0-83 | 0-11 | 0:86 | 0-46 | 0-42 | 0-04 | 0-44
21. Lagos 5 — — | 0-85 | 0-78 | 0-07 | 0-86 | 0:46 | 0-42 | 0-04 | 0-44

* Relation of the shoulder width to the length of pronotum.
¢ Relation of the length of hind femur to that of the elytron.

When studying the first two lines of the table we see that individual figures for
specimens of both forms are highly variable. Owing to this variability there is no
interval between a row of figures for danica and that for migratoria, which partly
overlap each other. This accounts for the fact that a curve representing the
variability of the pronotal proportion (fig. 2) in both forms together has only one
maximum ; it seems to indicate that, so far as the shape of the pronotum is concerned,
there is no possibility of regarding migratoria and danica as different species, or
even as two distinct forms of the same species. The curve of variability of the
femoral proportion (fig. 3), on the contrary, has two separate maxima, as if the
material studied could be divided into two distinct groups, be these species or units
of lower taxonomic value.

As for the average figures for both forms, they are also quite distinct and even
not too near each other ; the maxima and minima are also more or less characteristic
for each form. It may be noted that danica is more variable than migratoria, the
extent of variation in it amounting to 35 per cent. of the average figure in the case
of the pronotal proportion and about 38 per cent. for the femoral; while the corre-
sponding variations in migratoria are only about 29 and 26 per cent. respectively.
The next interesting point is that specimens of danica from the Palaearctic region
are far more variable than those of extra-Palaearctic origin.

(3442) 7

140 B. P. UVAROV.

All subsequent lines in the table (from the fifth downwards) give figures each for

a number of specimens from one locality and taken mostly at the same time, without
any selection, and regardless of the forms to which they belonged, in order to obtain
an impression as to proportion of both forms and of the extension and direction of
variability in each lot separately. We shall have to deal with these lines in more

ri)
65
60
55
50
a”
e 4s
iJ
E
‘3 40
i]
=
~ |
Cae te ea
P]
eee
¢ 25
1 he ee
ale
MBA
a on
ACE
Sr Pps ay Te a ee a ee
Be SoS au Goce wears ee eee
ee ee ee ae ee
Scan Wed 40) 2 Ome eco

Pronctar proporttawm.

Fig. 2. Diagram showing range of variation in the pronotal proportion in 358 specimens of
Locusta migratoria, L.

detail afterwards, but it may be noted that the range of variation is rather different
not only in series of different origin, but also in lots taken at the same place in two
successive years, 7.¢., from swarms which may be regarded as two _ successive
generations. Examples of this kind will be found in lines 5 and 6, and 7 and 8,
of Table i.

If we summarise the results of this attempt at a statistical study of themorphological
interrelations between migratoria and danica, they seem to be rather contradictory,

AS REVISION, OF THE. GENUS. LOCUSTA, EL. 141

or, anyway, do not help much towards a definite solution of the question whether
these two forms are really distinct or not.

— - Ww ~~ j=2) a wy ~ n = oO

x~ > =n r sr RHR & in an KR Pr)

Or ee eA ies heh hed: est) es. wie fe

Be 4 1 t ‘ f] 1 i] i] f] 4 (
o = § <a x an

Selsey) cee et ee ees ta ey) ae ae

APO OF Baie Gn (Oy Ve as Ss Si os: WS

Semorak proportior

Tig. 3. Diagram showing range of variation of the femoral proportion in 358 specimens of
Locusta migratoria, L.

Characters drawn from the Genitalia.

The late Dr. N. Adelung, of the Petrograd Zoological Museum, who tried long
ago to solve the problem of migratoria and danica, suggested to me, when I began
to work at it, that the anatomy of the genitalia, especially those of the males, might
give definite proof of the specific difference of these insects, which he himself believed
to be distinct. He even prepared some rough sketches of the male genitalia of
both forms, which, in his opinion, showed that they may be separated without much
dithiculty by the shape of the penis. He gave up his work, but handed over to me
his sketches and photographs. After a careful examination of these, and a com-
parison with several good preparations, I am fully convinced that Dr. Adelung’s
conclusion is not right ; since the seeming difference between the penis in migratoria
and danica in his preparations (so far as I know, he dissected only one specimen of
each form) depends entirely on the fact that he studied the whole genital apparatus
without dissecting it, and the somewhat different shapes shown in two of his
drawings are merely the result of covering tissues having been more completely
removed in one case than in the other.

142 B. P, UVAROV.

My studies show, on the contrary, that the male genitalia of danica and migratoria,
which are composed of several very complicated pieces (see fig. 7, A, B, C, D) rare
rather constant in the shape of the different parts, and though slight variations do
occur, these are in no way connected with the external differences, and afford no
reliable criterion for separating the two forms.

Colour Characters in Adults and Larvae.

In regard to the general coloration, as well as the pattern of the different parts,
adults of both danica and migratoria are rather variable, and no definite colour
characters can be given for separating them. Generally speaking, however, the
extent of the colour variability in migratoria is far less than in danica, and there
is a well-marked tendency in the former to a paler general coloration and less defined
markings ; especially noteworthy is the fact that the hind tibiae in migratoria are
never red, save quite exceptionally.

The coloration of danica is generally far more variable, with the evident prevalence
of bright green forms; dark brown, even almost black forms occur also not
infrequently, while the pattern is very variable but always well pronounced. The
hind tibiae in danica are very often red, but this character is far from being constant,
since often quite typical specimens have the hind tibiae pale or greenish.

While, as above stated, the coloration of the adults is of but very little use for
separating the two forms, it is quite otherwise in the case of larval stages, in which
there is a very striking difference. In fact, it is almost impossible to find any definite
type of coloration of the larvae of danica, which vary enormously ; uniformly green
forms are most common, but fawn, grey, brown, and even black ones may be met
with together. Quite the opposite is the case in migratoria, in which each larval stage
exhibits quite constant colour characters. Their coloration presents a combination of
black and orange-red (or yellow), the earlier stages being almost entirely black,
while orange, or yellow, appears first in the third stage, extending gradually after
each subsequent moult.* There is, of course, a certain variability in the shade of
the colours, but as a rule larvae of migratoria of the same stage are all practically
identically coloured ; and it is important to note that this type of coloration never
occurs in the larvae of danica in spite of the wide range of variation in the latter.

Sexual Dimorphism.

There is a well-pronounced sexual dimorphism of the general dimensions in
danica, the males (adults) being distinctly smaller than the females ; average figures
of the length of the body, elytra, etc., for males differ from corresponding figures for
females by about 20 per cent. of the latter, so that one may separate the sexes by the
size only, provided that the specimens are all taken at the same place and time.
As for the proportions between the dimensions of the different parts of the body,
they are not connected with the absolute dimensions and remain the same in
both sexes.

In migratoria no such apparent difference in size between the sexes is observed,
the males being on the average smaller than the females by only some 4 per cent. of
the figures for the latter ; and males of larger size are indistinguishable from females
so far as the dimensions are concerned. Since the absolute dimensions in both
migratoria and danica, especially in the latter, are rather variable, they are of very
little value for separating these forms.

* I do not propose to give a detailed description of the larval stages of migvatoria, since this
has been done more than once by different authors; very good descriptions and fairly good
coloured figures have been given by H.C. Pratt (Dept. of Agric. Feder. Malay States, Bull.
No. 24, 1915). j

A REVISION OF THE GENUS LOCUSTA, L. 143

Another case of sexual dimorphism is observed in the adults of migratoria during the ,
period of copulation and concerns the general coloration.* The males gradually |
assume a bright yellowish general coloration, most pronounced on the pronotum,
while the females become more dull coloured, the pronotum turning to dark brown.
These changes doubtless depend on some physiological processes connected with the
maturation of the sex product$. As for danica, it is not yet known whether it
exhibits any change of coloration during adult life.

Biology.

While danica seems to have no marked preference for any particular type of
locality for its permanent habitation, except vast waterless tracts and_ forests,
nuigratoria, on the contrary, has strictly defined permanent breeding grounds. In
Russia, where much attention has been paid by entomologists to ‘the biology of
migratoria, its breeding grounds are all confined to the basins of the Caspian and
Aral Seas and of Lake Balkhash, and more definitely to the deltas of the rivers
discharging into them, namely, the Volga, Ural, Kuma, Terek, Arax, Syr-Darya,
Amu-Darya, Ili, etc. These deltas, as a rule, extend over vast areas irrigated by
numberless channels which change their course almost every year, some of them
forming temporary pools and small lakes. The shores of these channels and even
the beds of the shallower ones, as well as all the less elevated portions of land
separating the channels from each other, are covered with a dense growth of the
gigantic cane, or reed grass (Phragmites communis, Trin.), sometimes 10-15 feet high,
which form almost impenetrable jungles extending over hundreds of square miles.
These reed-beds, however, are not uninterrupted, since many of the islands between
the channels are more elevated above the level of the water than is suitable for the
growth of the reeds, which require a very damp soil. The soil of such islands
mostly contains a large proportion of sand, and a peculiar flora of low, not very
dense, mesophilous and xerophilous grasses covers them. It is in such localities
that migratoria lays its eggs, as a rule,t while its larval swarms wander all over the
valley, penetrating through the reed-beds, and even swimming across the streams.
The leaves of the reeds, which contain a very large percentage of silica, represent
the most preferred food of the larvae. The climatic conditions in these reed-beds
are very peculiar and differ very much from those prevailing in the adjoining steppes
and deserts ; since the waters of the river are there spread over a vast surface, the
evaporation is very extensive, and the damp, hot air amidst the reeds at midday
vividly recalls the tropics.

All the sufficiently investigated breeding grounds of migratoria in Russia are of
the character described, and nowhere except in these grounds does this insect live
and breed permanently. Naturally the locusts are not always equally numerous
in the breeding regions, and in the years of minimum development only a few swarms
or even but single “individuals may be found; during the next few years their number
increases, the swarms become more and more dense, and a maximum is in this
way attained.

The newly-hatched larvae collect in small groups, each of which is, as a rule,
formed by larvae hatched from one egg-mass. These groups soon begin to move
irregularly to and fro ; if two groups meet they form one larger group and in this way
larval swarms are formed, and their movements become more and more regular.
Most authors regard want of food as the direct cause of these movements, but this
explanation is entirely wrong, at least so far as concerns Locusta migratoria and

* This fact was first observed by my assistant, Mr. Th. Gliniuk, in 1912, in Stavropol province,
and was recorded by me in the paper: “‘ The Fight against Locusts in the Government of Stav ropol
in the years 1907—-1912,’’ St. Petersburg, 1913. (in Russian).

+ The matter is really more complicated than this, since the selection of spots for oviposition
is usually closely connected with the more minute character of the soil and vegetation.

144 B. P. UVAROV.

Dociostaurus maroccanus, Thunbg., both of which I have studied for several years.
It is true that when the vegetation is very rich, the rapidity and extent of movement
of larval swarms is less than in the case of a sparse vegetation, but larvae wz/l move,
however densely overgrown with their most favourable food-plants the place may
be. Moreover, in both the species referred to, the larvae do not feed during move-
ment, save in some exceptional cases, as, for instanee, when a swarm has previously
wandered for a long time over barren ground and then comes across a field of corn
or other rich vegetation. My observations, though far from being complete, leave
no doubt that a factor of utmost importance in the movement of larval swarms
is temperature, and the following outline of the daily regime of a swarm will help
to explain this. .

It is a well-known fact that larval swarms do not move at all during the night,
which they pass on plants in a semi-comatose state, caused doubtless by the low
temperature. The first rays of the rising sun bring the larvae back to active life,
and they begin at once to feed. The hotter the temperature grows, the more active
become the insects, and soon they one by one jump or crawl down to the ground,
where they continue to move about, now crawling, now jumping, as if feeling uneasy.
Each larva in doing so disturbs its fellows, which leads to still more rapid general
movement; this is at first entirely irregular, but sooner or later, through mutual
influence, a common direction of movement is found and the swarm begins its day’s
march. So far as our observations in the Northern Caucasus go, the average
temperature causing the first movements of the larvae lies near 13-15° C. In
ordinary circumstances, ?.e., provided the sun shines all day and no unusual changes
of weather occur, swarms keep moving the whole day, and as a rule do not feed
when they move.* In the afternoon, when the temperature begins to fall, the
velocity of the movement decreases until, in the evening, the swarm stops; it is
very interesting to note that this evening cessation of movement occurs when the
temperature is again near 13-15° C. One by one the larvae crawl up plants and
begin to take their evening meal. There is no doubt that swarms do not choose
their resting places, but simply stop where they are overtaken by the critical
temperature. I have seen many swarms which have stopped for the night at entirely
barren spots just after they have crossed a strip with abundant vegetation, or only
a few yards before reaching it.

Such is the behaviour of larval swarms of migratoria under ordinary conditions
of weather, and all my observations lead me to the definite conclusion that their
movement has nothing to do with hunger and depends entirely on thermotropism
(probably negative), and on another, as yet little known, tropism which is displayed
by the tendency of each larva to repeat the movements of its nearest fellows and to
move in the same direction as they do.t+

The behaviour of larval swarms under abnormal conditions of weather lends
further support to this conclusion. If the day is colder than usual and the tempera-
ture does not rise above 15° C., the swarm remains and feeds all day where it
spent the night. If the temperature falls during a normally hot day, the swarm
stops long before its usual time. Especially interesting and instructive is the
following example: if a swarm is moving at the temperature in sunshine not much
above the critical point and the sun is temporarily hidden behind a cloud, so that
the temperature falls below that point, the swarm stops, or in any case its movement

* I intentionally avoid discussing here the question of the direction of movements and its
causes, since it is very complicated, and besides, it has no direct connection with the chief problem
we are now investigating ; one point, however, is worth mentioning : that the direction of move-
ment has, as a rule, nothing to do with the looking for food, since, for one thing, larvae do not
feed during the day, and they often move from fertile spots into entirely barren places.

+ This kind of tropism is not uncommon among other insects, and even vertebrates ; a herd
of pee will afford excellent examples of it.

A REVISION OF THE GENUS LOCUSTA, L. 145

becomes slower and less defined ; as soon as the cloud has passed and the sun shines
again, the swarms starts its march afresh ; even small clouds hiding the sun just
for a few minutes produce the same effect.

There is likewise an upper limit of temperature that causes the cessation of
movement ; and on very hot, close days swarms often stop their march during
midday ; in that case, however, the larvae do not climb plants to feed but collect
in close clusters under the plants, evidently hiding from the direct rays of the sun.
This being a more rare phenomenon, I have no precise data as to the exact tem-
perature at which it occurs, but anyhow it gives additional support to my theory
that the movements of larval swarms depend primarily on thermotropism.*

After the final moult the wanderings of swarms on foot naturally cease, though
when the majority of a swarm is in the last larval stage and only single individuals
become winged, the latter may often be seen crawling and jumping with the swarm.

A few days after the last moult, newly-winged insects are incapable of long flight,
their elytra and wings being not yet hard enough. This period is passed by swarms
in the same spot, devouring vast quantities of food, and it is a very favourable
(and also the last) opportunity for their destruction by spraying.

When locusts are fit for flight, single individuals begin to take wing and fly for
a short distance, often circling above the still sitting swarm. Whenever a locust
flies near enough to another that is at rest, the latter is disturbed and often takes
wing and flies in the same direction ; this is again a manifestation of the same tropism
which causes the movement of larval swarms. The larger the number of individuals
with fully developed wings, the more often do they take these short flights, and
the more other locusts join them, disturbing yet others when circling above the
swarm. It is easy to understand that this must necessarily result, sooner or later,
in the whole swarm taking wing. During the first movements no definite direction
of flight is apparent, but since each individual tries to follow its nearest fellow, a
common direction of flight must necessarily result. During the first few days these
flights are rather irregular, and swarms do not assume a definite direction, but simply
circle above their breeding grounds. If two swarms meet, they mix together, and
so the swarm gradually grows larger and larger. The larger the swarms grow, the
more regular and the longer become their flights, and at last the time comes when they
assume a definite direction, and the insects take leave of their breeding region
altogether ; then only few scattered swarms remain where just a day or two before
locusts were numberless. In fact this emigration from the breeding regions is often
so complete that only single individuals are left behind, and those prove to be nearly
all parasitised by the larvae of Sarcophagid flies, or by red mites (Tvombidium).

What is the cause of this emigration? The generally accepted theory is that
locusts migrate from want of food. I have already proved that this is not the case
in the larval swarms, and as for flyers, the very idea of locusts being compelled to
emigrate from breeding regions by the lack of food could never occur to anyone who
has seen these vasts areas overgrown with luxurious vegetation of a kind most
acceptable to locusts. In fact, swarms always leave behind them immense feeding
grounds and emigrate sometimes to almost vegetationless deserts, which, as a rule,
adjoin the permanent breeding areas of migratoria.

* The habits of the larval swarms of migratoria are well known to those concerned with
locust control in Russia, and as the insects are now almost exclusively destroyed by spraying
with arsenical insecticides, which are of use only when the actual food of the larvae is poisoned
before feeding, no spraying is done during day, when swarms are in movement. The daily work
is usually divided into two periods, and spraying is done in the early morning, before the larvae
begin to go down from the plants, and in the evening; the evening work begins before the
swarms stop for the night, since it is always easy to reckon where a certain swarm will stop,
and the spraying is continued till it is quite dark; this evening spraying is the more effective.
In cool weather, when swarms do not move, work goes on all day continuously.

146 B. P. UVAROV.

Still more unacceptable becomes the theory of emigration being caused by lack
of food, if we turn our attention to certain physiological changes which locusts
undergo during the period of emigration. When dissecting individuals taken from
emigrating swarms, it will be found that by far the greater portion of the inner
cavity of the body is occupied by air-sacs, described long ago by American ento-
mologists in the Rocky Mountain locust, and occurring doubtless in all other
migrating species of locusts. These air-sacs are only temporary organs, reaching
their highest development at the period of emigration and disappearing towards the
end of that period, when the developing reproductive organs take their place.
During the emigration, however, the air-sacs are enormously large and all the other
internal organs are much compressed, including the stomach, thus rendering the
insect almost incapable of taking food, at any rate in large quantities. This
assumption, based upon anatomical facts, is supported also by field observations ;
for, in fact, the emigrating swarms, when they stop their flight, do not feed much,
though incidentally they may cause great damage by merely cutting the stems
of cultivated plants.

Further anatomical researches reveal also the fact that the fat-body is more
developed in insects just before and at the beginning of emigration, and is almost
exhausted towards the end of it ; probably locusts during this period live essentially
on the food reserves in the fat-body, being unable to take much vegetable nourish-
ment and consequently scarcity or even lack of food has nothing to do with the
emigration.

A Russian entomologist, K. N. Rossikov, called attention to another possible
explanation of the emigration of migratoria from its breeding grounds. He believed
it to be an immediate result of the activity of the parasites of the adult locusts,
i.e., Sarcophagid larvae and red mites ; he believed that the parasitised individuals
become restless and try to get rid of their parasites by flight. If this were so, the
result would be that only parasitised individuals would emigrate and all the non-
parasitised would remain behind in the breeding grounds, though actually just the
opposite is observed. This theory, therefore, is as groundless as the previous one.

One more theory is that emigration might be regarded as a tendency of the
species to avoid overpopulation of a breeding region and to find new suitable breeding
grounds. As for the possibility of overpopulation of breeding regions of migraioria,
this idea is simply absurd, since these regions are vast enough to harbour many hundred
times more locust swarms than there are in years of maximal development.

To investigate the presumption that swarms are emigrating to look for new
breeding grounds we must see what is the fate of swarms after they have left their
permanent breeding regions.

As far as is known at present, a swarm of emigrating locusts usually covers a
very long distance at one flight ; if sometimes it settles down on its way (and this
is often caused by unfavourable conditions of weather), it soon resumes its flight
again. I will not discuss here the question of the direction of the flight and its
probable causes, since but very little is known about it. One fact, however, is firmly
established and is of great importance for our immediate purpose : it is that in the
majority of cases the swarms maintain throughout the same more or less defined
direction which they assumed when starting ; of course, a strong wind or other
incidental circumstances may to a certain extent alter this direction.

This straight flight, aimless and causeless as it seems, does not last long, though
a swarm may cover during it very long distances, the velocity of flight being far
greater than might be expected. j

Sooner or later, the regularity of the flight seems to be lost ; swarms begin now
to settle down, then take wing again and circle about ; they begin also to eat more,
since their air-sacs have grown smaller and the fat-body is also exhausted. I believe,

A REVISION OF THE GENUS LOCUSTA, L. 147

therefore, that the cessation of flight is caused entirely by these two purely physiological
phenomena, and it is obvious that no possibility exists for swarms to look for
suitable new breeding grounds ; they merely settle down wherever they are compelled
to do so by their physiological condition, quite irrespective of the character of the
locality. This may occur accidentally near some suitable spot, and there is also
the probability that the swarm would find one during the irregular circular flights
which take place at the end of the emigration and precede pairing and oviposition ;
but the extent of these flights is rather limited and this probability is correspondingly
small. Theoretically it is far more probable that the cessation of emigration would
become necessary in a locality quite unlike the normal breeding grounds.

A typical and very convincing example of this kind was observed in Stavropol
province in the autumn of 1912, when numerous large swarms of migratoria
emigrating from the breeding area at the mouth of the river Terek invaded that
province. One or two of these swarms settled down in the lower portion of the
valley of the river Kuma, which is itself a breeding region of the same locust, but
was in that particular year free from the local swarms; the Terek swarms con-
sequently found there most favourable conditions and oviposition took place in the
normal manner. Practically all the other invading swarms stopped their flight
in the steppe adjoining the middle course of the Kuma, some of them on the very
border of the valley. Now this part of the valley presents some very suitable breeding
grounds, which have often played an important part as the source of invasions in
Stavropol province. Several of the swarms visited these grounds more than once during
their circular flights, which are often supposed to serve the purpose of finding
suitable places for oviposition. Ultimately, however, only a small number of
scattered locusts oviposited there, while all the swarms deposited their egg-masses
in the dry steppe, where the conditions of soil and vegetation are entirely different
from those in normal breeding grounds. It is especially interesting to note that
some of the eggs were laid on a portion of the steppe sloping towards the valley of
the river, 7.e., in the closest proximity to the above-mentioned suitable area.

This latter fact and a study of the general conditions under which oviposition
took place clearly show that nothing in ‘the least like a conscious (or instinctive—
the exact word does not matter in this case) choice of suitable places by swarms can
be assumed. There is, however, one exception: when a swarm settles down for
oviposition, and the females, after several attempts to penetrate the soil, find it too
hard, they become restless, take wing again, and after a few rounds settle down at
another spot. Thus we must conclude that oviposition takes place whenever the
majority of females are ready for it, and quite irrespective of the suitability of the
conditions for the next generation, provided that oviposition is physically possible.

In conclusion, the theory that emigration has as its aim the finding of new
breeding places is also groundless, and there is at present no possibility of explaining
the emigration by any causes except phy siological ones: the development of the
air-sacs compels the insects to fly, and this impulse is strengthened by their gregarious-
ness, that is by some kind of tropism which makes each individual keep close to its
fellows and follow their movements. Later on we shall see what is the biological
meaning of the emigration.

Such is, briefly, the life-cycle of migratoria. The biology of the larvae and adults
of danica is only very insufficiently known,* but what is known shows that their
behaviour is entirely different from that of migratoria. The chief biological feature
of the latter in both larval and adult stages—gregariousness—is quite absent in
damca. This is especially striking in the larvae ; if a wandering swarm of migratoria

* This is directly due to the fact that most entomologists have regarded danica as distinct
from migratoria and as being an entirely harmless species, so that the study of its habits has been
neglected.

148 B. P. UVAROV.

larvae comes across a solitary specimen of the same form, the latter immediately
joins the movement, but when a larva of danica is overtaken by a swarm of migratoria,
it tries to escape by leaps as quickly asit can. At the same time, the larvae of danica
seem to possess thermotropism of the same kind as that exhibited by migratoria,
their time of feeding being restricted to the evening and early morning, while
during the day they are probably also on the move, though I have no reliable
observations on this point. As for the adults of danica, the only point of their biology
that we know for certain is that they do not form swarms and hardly migrate at all ;
a study of their behaviour, as well as of their anatomy (air-sacs) is of the greatest
importance for the solution of the whole problem.

Some very interesting indications of further biological differences between
danica and migratoria have been obtained at the Turkestan Entomological Station
by V. Plotnikov in his breeding experiments.* This entomologist obtained from
typical individuals of danica, kept in the laboratory, a second generation of larvae
in only 16-30 days after oviposition, instead of in the following spring as is usually
the case with migratoria. In one particular experiment even three generations
were bred in one year. That this unusually short period of hatching was not due to
the unnatural conditions of the experiment is shown by the fact that eggs laid in
the same laboratory by individuals of migratoria did not hatch before the following
spring. Dissections of eggs showed that the development of the embryo begins
in eggs of both migratoria and danica shortly after oviposition ; but in the case of
migratoria, when the embryo reaches a rather advanced stage, development is
suspended for several months, corresponding to the period of hibernation, though
in the laboratory there is no change in the conditions to account for this. At the same
time and under exactly the same conditions, the embryos in eggs of danica develop
without any interruption. These experiments suggest an explanation of the fact
that, while the larval stages of migratoria may be found only in spring and the
beginning of summer and the adults during the summer and autumn, there is no such
strict regularity about the occurrence of the stages in danica, though many eggs of
this form probably hibernate as well.

Field Observations on the Transformation of mgvatoria into danica.

During the great invasion of locust swarms which occurred in the Stavropol
province in the autumn of 1912 (see p. 147), I used the opportunity for studying,
from the systematist’s standpoint, as large a series of specimens as possible. All
the insects collected, which were taken from the swarms without any selection and
amounted to many hundreds, proved to be quite typical migratoria. Nothing in the
least like danica was observed in field, either by myself or by my assistants, whom I had
previously instructed to look out for all aberrant forms and who knew danica perfectly
well; the number of individuals thus studied without collecting them is difficult to
estimate, but it doubtless amounted to many thousands. I believe, therefore, that
I am right in assuming that the swarms consisted purely of migratoria, and that
danica, or even intermediate forms, were entirely absent. The measurements of
the specimens from those swarms are given in the fifth line of Table I (p. 139),
and the following conclusions may be drawn from them: the specimens are rather
uniform, the extent of their variability (0-16) being less than the average for migra-
toria (0-18—see line 1) ; the average figures for the pronotal (0-79) and the femoral
(0-45) proportions are extremely near to the average for migratoria (0-80 and 0-46,
respectively). The colour characters, though not very reliable, were very constant,
which is not the case in danica. If we consider also that the locusts kept in close
swarms which had no tendency to disperse, we must conclude that the swarms were
formed exclusively by typical individuals of migratoria.

* Report on the work of the Turkestan Entomological Station in 1912, 1913, 1914, and part of
1915; pp. 28, 55-59; Tashkent, 1915 (in Russian) ; see also Rev. Appl. Entom., iv, p. 211.

A REVISION OF THE GENUS LOCUSTA, L. 149

According to the routine of the control work adopted in Russia, all the swarms
were closely watched during their wanderings by the trained staff, and all the spots
where oviposition took place were marked out and also noted on the maps. Owing
to this procedure there was no doubt that in the following spring we had to conduct
the destruction work against the immediate progeny of those swarms. As soon as
the larvae in 1913 reached their third stage, when differences between migratoria
and danica are more apparent than in the earlier stages, it became evident that
although the bulk of the larval swarms consisted of migratoria, there were many
individuals which were certainly dantca, these being different in coloration and showing
a tendency to desert the swarms. In spite of the intensive control measures, several
small swarms escaped destruction and attained their final moult; these adult
specimens proved to be rather different from those of their parental swarms (see
line 6 of the table, p. 139). A rather large admixture of typical danica was very
obvious, but still more numerous were specimens of an intermediate character which
could not be identified either with danica or with migratoria ; the bulk of the insects,
however, might have been referred to migratoria, but showed an obvious inclination
towards danica, in fact they had the pronotum more compressed laterally, the median
keel more raised, the elytra longer and the femora relatively shorter, than in the
specimens from the parental swarms; their coloration was also more variable.
The proportions are especially instructive when compared with those for the swarms
of 1912; in studying these figures one may see that the extent of variation was far
larger than in 1912, and the average figures also changed in the direction of danica.
The swarms were not so dense as in 1912, and the individual insects showed obviously
less developed gregarious habits ; numerous single individuals of danica were scattered
all over the steppe, without any connection with the swarms; the latter did not
undertake any migrations and gradually dispersed.

Similar, though less striking examples are given in lines 7 and 8, 9 and 10 of
the table, in compiling which precautions were also taken to obtain the series most
likely to represent two successive generations. Unfortunately, I could not secure
reliable examples of more than two such generations from one spot.

Breeding Experiments by V. Piotnikov.

Though field observations like those described above are of great value, they
have the disadvantage of not affording absolute proof, and the only way to obtain
this is by breeding experiments. Such experiments have been undertaken by my
friend V. Plotnikov, in Tashkent, and, though conducted on a very moderate scale,
have yielded some most interesting and valuable results. Since all the actual
specimens from these experiments were given to me by V. Plotnikov, and are before
me now, I am able to give a little more detailed account of the results than were
recorded in his original communication.

In the summer of 1913 several specimens of both sexes of very typical danica
were isolated in cages, in which copulation and oviposition took place; the eggs
hatched without hibernation, as is not uncommon with danica, but so far as we know
never occurs in migratoria. The description of the larvae and adults bred from them
is given by V. Plotnikov, as follows :—

“The larvae had in the first stage a dark grey coloration, and not black as in
migratoria. In later stages they acquired various colorations—uniformly green,
dark grey or brownish—but a number of them had the typical colouring of migratoria,
namely, a general reddish brown colour (sometimes greenish), with velvety black
stripes (broad or narrow) along the sides of the pronotal keel and black stripes on
the sides of the abdomen. The adults presented no characters typical of danice ;
the profile of the pronotal keel was usually straight, sometimes even concave. The
males were, however, smaller than the females.”’

150 B. P. UVAROV.

After studying the specimens, I can only confirm Plotnikov’s statement that while
the parents are all very typical danica, save that not all of them have the hind tibiae
red (which character is not quite constant in that form), their direct offspring are
on the contrary all well-defined migratoria, though a few of them have the tibiae
red, as is sometimes the case in this form. One of the parents and one of the
off-spring are figured above (fie, GAS BC eeeiD):

Another experiment is described by V. Plotnikov, as follows :—

“Tn 1914 I bred from egg-masses sent from Amu-Darya district* P. migratorius,
and from egg-masses deposited by these insects I bred in the spring of 1915 again
migratorius. On the 19th June I found in the soil of the breeding cage, where these
individuals (now mature) used to live, five egg-masses, which I transplanted carefully
into the soil of another cage; there, on the 6th August, 7.e., more than 48 days
after the oviposition, a single larva hatched, a female of dark grey coloration ; the
rest of eggs in the egg-masses remained with an halt-developed (hibernating)
embryo. When in its second stage, this larva acquired a green coloration, which it
retained till the fifth (final) stage. The profile of its pronotal keel was convex. . . .
The adult insect retained the convex keel of the pronotum; its body was green,
the elytra light brown, and its hind tibiae turned red.”’

This specimen is before me now, and I can only confirm V. Plotnikov’s opinion
that it represents the most typical danica. As for its actual parents they possess
all the essential characters of migratoria very well defined, and no one could hesitate
to identify them with that form. Unfortunately the experiments were discontinued
upon Plotnikov’s joining the army.

V. Plotnikov’s conclusion from his experiments is as follows: pat as
impossible to separate P. migratorius and P. danicus by any characters ; characters
of danicus (including its capacity to produce a second generation) are expressed in
the latter species more strongly. It is possible to suppose that this species is now
in the process of splitting off from the primitive species, P. migratorius.”

My own conclusions differ somewhat from this, but I shall come to them later
on. All that I shall point out now is that these experiments prove finally the
possibility of the actual breeding of migratoria from danica and vice versa; My
field observations on the same subject give evidence that it may occur not under
laboratory conditions only, but in nature as well. On the other hand, we must
not forget the numerous differences between them, especially the biological ones,
which prevent us from regarding danica as a mere synonym of migratoria. It is
evident that they must be regarded as two different forms of the same species without,
in the meantime, any more precise definition of their systematic value.

Locusta migratoricides, Rch. & Frm.

This insect was described from specimens from Abyssinia; later on, Saussure
and other authors recorded it from many tropical localities. In its morphological
features it is very much like migratoria, while its difference from danica is far more
marked than in the latter. From migratoria it differs only in the following
characters :—The pronotum (fig. 4) is still more constricted before the middle ; its
median keel very low, often concave in profile ; fore margin almost straight ; hind
margin very widely rounded ; the shoulder width almost equal to the length of the
pronotum, the average pronotal proportion being 0-86, while it is 0-80 in migratoria ;
the elytra relatively longer and the hind femora shorter, which results in the femoral
proportion being on the average 0-44, as against 0-46 in migratoria (see Table’ J;
lines 20 & 21).

* A permanent breeding region of L. migratoria.—B.U.

A REVISION OF THE GENUS LOCUSTA, L. L5i

It is quite obvious from this definition that migratorioides presents no new features
as compared with migratoria, but it seems to be in all the chief characters
merely a further modification of migratoria in the direction opposite to danica.
Unfortunately the material of mzgratoriotdes now at my disposal is rather scanty,
and the extent of its variability remains uncertain. Still, there is in the British
Museum one female taken at Sarkwalla, Northern Territories, Gold Coast, 4—7.xi.1915
(Dr. J. J. Simpson), which is in all respects intermediate between migratoria and
migratorioides.

BR felling

5 2 . ° rie ne A
Fig. 4. Locusta migratoria, ph. migratorioides, Rch. & Frm., G from Lagos: A, head and pronotum
from above ; B, pronotum, side view ( 3).

The study of the male genitalia of migratorioides reveals no difference whatever
from the structure observed in danica and migratoria (fig. 7).

As for the coloration of the larval stages, we have a good description of them
given by H. C. Pratt (/.c.) and from it, as well as from the study of the actual
specimens sent by Mr. Pratt to me in 1913, I was unable to find any reliable difference
between them and the larvae of migratoria.

On the other hand, Mr. Pratt says in his paper that the coloration of the larvae
is not very constant and the green individuals occur alongside with the typically
coloured ones. The figures of the green and black adults given in his paper (J.c.,
pl. XIV) certainly represent danica. This is, then, an indication that the same inter-
relation exists between migratorioides and danica as has been proved positively for
migratoria and danica. A confirmation of the same fact I received not long ago,
when several specimens of locusts were sent from North Borneo to the Imperial
Bureau of Entomology for identification. They all proved to be migratorioides, but
as the lot was rather small, the Bureau asked for more material, and after several
months a new collection arrived, with a note that the specimens were taken singly
at the same spot as the swarms from which the first lot had been collected, and
represented the actual offspring of those swarms. All specimens in this second
lot are quite typical danica.

One more example of the transformation of migratorioides into danica I have found
in Dr. La Baume’s paper on the African locusts.* In a reference to Dr. L. Sander’s
book} he discusses an invasion of locusts which took place at Misahdhe, Togo, in
December 1893, while in March 1894 new larval swarms appeared at the same
locality, which Dr. Sander believed to be the direct progeny of the December swarms.

* Die Afrikanischen Wanderheuschrecken.—Beih. zum Tropenpflanzer, xi, No. 2, 1910,
p. 82, footnote 22.

j Die Wanderheuschrecken und ihre Bekampfung in unseren Afrikanischen Kolonien.
Berlin, 1902.

152 B. P. UVAROV.

These latter consisted, as Dr. La Baume stated after a study of the specimens, of
migratorioides, while the March swarms, again according to his determination, were
composed of danica, which leads him to the conclusion that they could not possibly be
the direct progeny of the December swarms. I believe, however, that Dr. Sander
(who simply did not distinguish mzgratorioides from danica) was right, and that the
December swarms of migratoriotdes actually produced the March swarms of danica.

All these facts, of course, do not furnish us with absolute proof of the trans-
formation of migratorioides into danica ; but since such a transformation is firmly
established for migratoria and danica, I feel justified in assuming it to be true in this
case also.

Geographical Distribution.

Of the three forms here dealt with, danica has the most extensive range; in
fact, it is found throughout the whole of the Eastern hemisphere, except the coldest
regions beyond 60° northern and southern latitude, very high mountains* and vast
waterless deserts. As for the Western hemisphere, though many books state that
danica occurs in America, I know of no reliable evidence to this effect, and am fully
convinced that all the older records are due to mistakes, either in labelling the
specimens or in their identification.** Thus, F. Walker described Pachytylus brasi-
liensis, which is conspecific with danica, from a specimen in an unlabelled collection
which included insects not from Brazil only, but from other parts of the world also,
and the mistake in locality in this case is beyond doubt.

A form with such an enormous area of distribution may be expected to present
some geographical variations, and in fact, apart from the individual variability
which, as I have already stated, is very great in danica, some more constant varia-
tions, probably connected with geographical conditions, are also observed. Thus
Australian and New Zealand specimens are rather small on the average, with a
comparatively short pronotum and the wings slightly infumate; they have been
described by Saussure as a distinct species, Pachvtylus australis, Sauss. As, however,
no one has yet studied extensive series of individuals of Australian origin, the
constancy of these characters is not proved; on the other hand, specimens of the
same kind occur incidentally in other localities as well, and their distinctions from
the typical danica do not exceed the average extent of observed individual variability.
Again, as I have already pointed out (p. 139), there is a slight difference between
the individuals of danica from the Palaearctic region and those of tropical origin,
which also may depend on geographical conditions ; but in this case likewise more
extensive investigations are wanted.

The area of distribution of migratorioides, though not so extensive as that of

danica, still occupies the greater part of the latter, in fact the whole of it except
the Palaearctic region.

The latter region is the home of migratoria, which is however common in its
south-eastern parts only (especially in the basur of the Black, Caspian and Aral Seas
and that of Lake Balkhash), where its permanent breeding grounds are, while its
emigrating swarms penetrate much farther north and westwards, sometimes as far as
Finland and England. On the other hand, single individuals, which agree in all their
morphological characters with migratoria, occur far beyond the Palaearctic region
as well. Thus, Brunner v. W attenwyl recorded} migratoria from the Malay
Archipelago (Batjan and Borneo). I have myself seen a quite typical specimen of

* T have recorded (Revue Russe d’Entom., xiv, 1914, p. 232) this form from the Pamir upland
as high up as 11,000 ft. above sea-level ; there is in the British Museum a couple of specimens
taken at Giangtse, Tibet, at an altitude of 13,000 ft.

** Dr. J. Rehn, of Philadelphia, informed me recently that he is of the same opinion.

t Abh. Senkenberg. Naturf. Ges., xxiv, pp. 194, 196.

A REVISION OF THE GENUS LOCUSTA, L. i oy}

migratoria sent from Southern Celebes by Dr. Roepke to the Imperial Bureau of
Entomology, which had the following note attached: ‘‘ A Locustid from S. Celebes
(Pangka djene) appearing there in small swarms and causing damage to the natives’
plantations, such as rice, corn, etc.”’

The Theory of Phases.

Though the above recorded facts by no means exhaust the points to be considered
in connection with the problem of the interrelations of migratoria, danica and
migratorioides, they yet permit us to make an attempt to find out the best explanation
at present possible. The following theory seems to me to agree in a rather satis-
factory manner with all the facts known at present, though some modifications of
it may prove to be unavoidable when new data are available.

As a starting point, I take it as positively proved that the three forms
cannot be separated specifically and that they represent taxonomic units of lower
grade than the species, which must be called, according to the law of priority,
L. migratoria, L. They are, however, quite distinct from each other, though
connected by transitional forms.

What term, then, should be applied to them? They are certainly not mere
individual aberrations—as they are often assumed to be by other authors—since
they are rather constant in their average morphological characters and still more
so in their biology; nor can we call them subspecies, 7.e., geographical races, as
they are found together in the same locality; nor are they seasonal forms, since
the transformation of one of them into the other has evidently nothing to do with
season. The only more or less suitable term for them is “‘ morpha,” in the sense
proposed by A. P. Semenoy-Tjan-Shansky,* who proposed to apply this name to
such forms of a species that present a direct result of the immediate external
influences on the individual insect during its development, and which therefore do
not appear in succeeding generations if the original influence ceases. Under this
definition come seasonal forms, which may be obtained by the artificial application
to developing individuals of certain factors causing their appearance under natural
conditions ; the forms resulting from feeding larvae by some special food, etc. The
same term may be applied, according to the personal opinion of A. P. Semenov-
Tian-Shansky, expressed in his letters to me, to the case of the Locusta forms. It
seems to me, however, that the term “ morpha”’ is rather vague, and moreover
we are yet far from knowing whether the transformation of one form into the other
is due to some immediate external influence or to some yet unknown internal cause ;
I think, therefore, that the term “ phase ” (Latin phasa ; abbreviation—ph.) suggested
to me by Dr. G..A. K. Marshall is more appropriate, and its meaning will be mugiele
clear in the course of the explanation of my theory.

There is no doubt in my mind that migratorioides is the oldest form of the species,
since its morphological and colour characters are far more constant in comparison
with the more plastic migratoria, to say nothing of the extremely variable danica.
The permanent breeding regions of migratorioides have never yet been investigated ;
the only description of breeding places of this form in the Malay States given by
H.C. Pratt (l.c., pp. 6-7) must obviously be referred not to the permanent breeding
grounds, but merely to the places where the oviposition of emigrated swarms took
place. All we know at present concerning the permanent breeding areas of
migratortoides 1s based on the records of the occurrence of its swarms; and these
data enable us to state that the best conditions for the development of this form
seem to be present in tropical countries with a rather damp and hot climate, but
undoubtedly not in forests. Since, on the other hand, these breeding grounds seem
to be yet undiscovered, we may presume that they are also not in open, grassy iand,

* Die taxonomischen Grenzen der Art und ihrer Unterabteilungen. SBerlin, 1910.
(3442) M

154 B. P. UVAROV.

which is easily accessible and mostly cultivated or, anyhow, populated. I believe,
therefore, that permanent breeding grounds of muigratorioides are to be looked for
somewhere deep in the impenetrable jungles, overgrown with high grasses, reeds,
and such-like vegetation ; but even if I am mistaken in this supposition, it would
not affect my theory, which is based on the indubitable fact that the permanent
breeding of migratorioides is possible only in localities with certain natural conditions,
whatever those conditions may actually be.

Another well-known fact is that the development of migratorioides in its breeding
grounds does not go on always at the same rate, but that it is subject to a periodical
rise and fall, though the exact cause of the increase of locusts is entirely unknown.
When the increase is at its height, large swarms are formed, and their emigration
follows. Such emigrated swarms settle down and oviposit whenever they are
compelled to do so by purely physiological causes, and their progeny undergoes a
transformation into the solitary-living phase—danica. The very plastic, easily
adaptable, and in all respects more progressive danica must play an important part
in the extension of the range of the species, gradually but steadily populating new
regions. Being a product of a mutation arising partly from some unknown internal
cause and partly from outer (probably climatic) influences, danica is naturally subject
to sudden displays of atavism, which results in the transformation into the ancestral
phase migratorioides. We do not know yet whether this phenomenon can occur
spontaneously as a result of some internal physiological factor, but there is no doubt
that it is much favoured and often probably caused by the oviposition of danica
taking place under conditions like those of the permanent breeding grounds of
migratorioides. The gregariousness of the migratorioides phase is, of course, one of
the causes of a rapid increase in the number of individuals and swarms, and soon—
in the course of a few generations—the size of the swarms reaches the maximal
point, which is followed by emigration. In this way the dispersion of the species
goes on alternately by the gradual spreading of the danica phase and by the periodical
extensive emigrations of migratorioides. As a result, the species is now distributed
all over the Eastern hemisphere ; but, as we know, the distribution of migratorioides
is limited to tropical regions only, while danica goes over to the Palaearctic region
as well, where the swarming phase of the species is represented by migratoria. This
latter fact might be satisfactorily explained by the impossibility of finding in the
Palaearctic region the natural conditions exactly like those of the tropical breeding
grounds of migratorioides, chiefly in regard to a combination of heat and dampness.
The above-described (p. 143) reed-beds of Phragmites in the south-eastern part of
the Palaearctic region represent in all respects the nearest possible approach to
tropical conditions. This statement is strongly supported by the fact that the
fauna of these reed-beds includes two more Acridians of an undoubtedly tropical
origin; these are Gelastorrhinus sagitta, Uvar., and Oxya turanica, Uvar., both
described* from the valley of the Amu-Darya, in Transcaspia, and the former found
also on the River Kura, in Transcaucasia. Though very peculiar, and in the summer
recalling the tropics, the climatic conditions of these reed-beds are, of course, not
tropical, and their effect on the progeny of danica breeding there is not the same
as in the tropical breeding grounds of migratorioides: the reverse transformation
of danica into a swarming phase does not reach the phase of migratorioides, but stops
half-way at the migratoria-phase. This seeems to indicate that the transformation
is due primarily to the direct influence of external conditions, its extent being
proportional to changes in the latter, but only precise laboratory investigations can
help to clear up this complicated problem. It is interesting to recall here that
individuals of migratoria incidentally occur in tropical countries also (see p. 152),
and we may presume that their appearance is due to some abnormal conditions
of the development.

* Horae Soc. Entom. Ross, xl, No. 3, 1912.

A REVISION OF THE GENUS LOCUSTA, L. j eyo)

Little is known yet as to what happens in the breeding grounds after the
emigration,* except that the number of locusts drops suddenly to a minimum.
I presume that scattered swarms of the gregarious phase, as well as the progeny
of individuals of the solitary phase, cause the gradual increase in the number of
swarming individuals, and after a few years a new emigration occurs.

Thus, the periodicity of locust invasions is caused entirely by the wonderful
phenomenon of the transformation of a swarming locust into a solitary, harmless
grasshopper. Of course, the outline here sketched is necessarily rough, and the
actual proceedings are far more complicated, but the theory seems to me to be the
best possible in the circumstances.

The biological result of these phenomena is that the maintenance and dispersion
of the species is ensured in all circumstances: the swarming phases enable the
species to extend at one stroke its area of distribution to distant regions, and its
dispersion to the remotest islands is undoubtedly due to emigrating swarms; on
the other hand, the well protected and easily adaptable solitary phase secures a strong
footing in the countries thus reached, and under favourable conditions gives rise to
new emigrants ; the results achieved show that such an arrangement has been
extremely useful to the species. Even the most radical changes in the natural
conditions of the permanent breeding regions would result not in the extermination
of the species, but only in its transformation into the more adaptable danica phase.

An example of that kind occurred in Southern Russia. Though the now existing
permanent breeding regions are restricted, as I have described above (p. 143),
to the valleys of the rivers discharging into the Caspian and Aral Seas and Lake
Balkhash, the deltas of rivers emptying into the Black Sea (7.e., Kuban, Don, Dnieper,
Danube, etc.) also harboured not very long ago—up to the end of the eighties of the
last century—some permanent breeding grounds of migratoria. At present, however,
only the lower valley of the Danube is still a breeding region, while the valleys of
the other rivers of the basin of the Black Sea no longer serve that purpose. This is
easily explained by the fact that the valleys of the Don, Kuban and Dnieper were
during the end of the last century more or less cultivated or, at any rate, their
natural conditions were entirely changed by the persistent grazing of herds of cattle.
As a direct result of this the possibility of the transformation of the solitary phase
into the swarming one exists there no longer, and though the transformation takes
place incidentally, single specimens of migratoria being not uncommon, their numbers
do not increase, nor are swarms ever found.

The theory of phases suggests the theoretical possibility of the control of migratoria
by some measures directed not against the insect itself, but against certain natural
conditions existing in breeding regions which are the direct cause of the development
of the swarming phase. The above-quoted example in South Russia shows that
even comparatively slight cultivation of breeding regions leads to the desired changes ;
but the conditions necessary for the breeding of the swarming phase have not been
exactly studied, nor are the direction and extent of such changes known. The
first step, therefore, should be the most careful investigation of all existing, as well
as extinct, breeding regions, together with parallel breeding experiments under
laboratory conditions ; on the basis of results thus gained a system of theoretically
useful and practically possible measures for the conversion of breeding regions may
be outlined.

IV. LocustaNA PARDALINA, WALK., AND ITS PHASES.

My personal knowledge of this locust is limited to the study of preserved specimens,
especially of a large series sent to the Imperial Bureau of Entomology by Mr. J.C. Faure,

* The direct cause of this ignorance is that injurious insects, and locusts especially, are
studied only in the years of maximum development, and nobody cares about them in the minimum
years, when the clue to the whole locust problem is most likely to be found.

(3442) M 2

156 B. P. UVAROV.

of the Division of Entomology, Pretoria. Before proceeding to my own observations
on the morphology of the species, I will quote an extract from a letter from Mr.
Faure, dated 14th October 1920, which includes some very important and interesting
information on the question of the phases of L. pardalina.

‘My personal experience with the species began in the summer of 1914-15,
when scattered swarms began to appear shortly after the break-up of a prolonged
and very severe drought. Voetgangers (7.e., nymphs) of all stages and flyers occurred
together in loose swarms, and it was practically impossible to destroy them by the
usual method of poisoning. The swarms did not move in the usual compact forma-
tion, nor did they camp for the night in dense clusters. Many of the adults were
strikingly undersized, and a large percentage of both adults and voetgangers were
abnormally coloured. Only in swarms that approached the normal in density did
the typical orange and black colour of the voetgangers begin to show up.

« Although I did not realize the fact at the time, I was witnessing the transition
from the grasshopper to the swarm phase. Towards the winter, that is in May and
June 1915, the flyers began to move about in fairly definite loose swarms, and they
laid their eggs in compact deposits, with the result that large swarms of typical
swarm voetgangers hatched the following spring. We received no reports that
winter of swarms of flyers coming into the Union from the Kalahari or anywhere else,
and the outbreak in the period September to December 1915 was very severe in
the area in which the scattered locusts had been observed the previous summer.

“It was quite evident, therefore, that the invasion of September-December 1915
had arisen from locusts bred up within the borders of the Union. Formerly the
Kalahari Desert had been thought to be the chief source of our invasions of Locusta
pardalina. Now we are convinced that large outbreaks can and do arise within our
borders without the help of swarms coming in from the Kalahari. In the past,
huge swarms have undoubtedly come into the Union from the Kalahari, and no
doubt history may repeat itself in the future. But we no longer regard the Kalahari
as a sort of permanent breeding ground, and are now inclined to believe that it will
ordinarily only develop into a breeding ground if we allow swarms of flyers to escape
into it from the Union.

“Tn 1917 I again saw scattered locusts from February to April, and another
severe outbreak of voetgangers occurred the following spring and summer. In a
general way it was a repetition of what had occurred in 1915, and realizing what was
going on, I was better able to make observations.

‘ Locusta pardalina does not merely occur in scattered swarms and in compact
swarms—it also lives as a grasshopper, 7.¢., single specimens have often been collected
miles away from the nearest swarm and in seasons when no swarms have been known
to exist anywhere in the country. I have good reasons for believing that the species
is probably never entirely absent from certain parts of the Union. During the
past five years I have often searched for specimens during the intervals between the
occurrence of swarms, and in practically every case I have succeeded in capturing
two or three at least in say an hour’s walk on the veld. Of course one should not
expect to find them late in the winter or during a bad drought.

‘The specimens captured singly almost always have the colours of the grass-
hopper phase, and they are as a rule a good deal smaller than swarm forms. Further,
I have frequently taken last-stage nymphs and newly-fledged adults living the life
of single grasshoppers. Although I have not been able to make a careful study
of the specific characters of these single living forms, Iam quite satisfied in my own
mind that they are identical with the swarm forms. Again, these single forms may
occur in the district or on the farm on which swarms are present, and I have on
several occasions seen a few individuals with abnormal colours amongst a swarm

A REVISION OF THE GENUS LOCUSTA, L. VO7

of typical swarm-form voetgangers. The only conclusion I could come to was that
these stray forms have been picked up by the swarm. I have also found grass-
hopper-phase adults in a swarm of swarm-phase flyers, and have seen a very small
green-marked male in copulation with a large typical female.

“These single forms of L. pardalina can readily be distinguished in the field
from our other veld grasshoppers (1) by the fact that they have milky-white glistening
underwings, and (2) by their peculiar manner of flight. They almost always soar
upwards and then dip and swerve before settling down.

“From the swarm-phase the grasshopper-phase of L. pardalina differs chiefly
in size and in colour. As regards colour, the single forms are remarkable for their
great variability, and it would be quite an undertaking to describe in detail all the
shades of colour they exhibit. Usually there is a striking protective resemblance.
Where there is plenty of green grass both voetgangers and flyers may be almost entirely
green, or at least partly green. Where the veld is only sparsely covered with grass
and bushes, they resemble the colour of the soil more or less. In parts of the Karroo,
notably Beaufort West and Prince Albert, there are patches of gravelly soil varying
from slaty-blue to almost black. In 1917 I was greatly surprised to find a very
striking tendency amongst the scattered veotgangers to vary in colour from place
to place more or less in accordance with the colour of the soil. When the progeny
of these scattered locusts appeared on the same farms in swarms six months later
there was no trace of such a protective resemblance; they all wore the King’s
tegulation swarm uniform !

“ As far as size goes, the single-living forms are generally considerably smaller than
the swarm forms. This is especially true of the males. Some of the males are so
small that one can scarcely believe that they belong to the same species as the
swarm males.

“When it occurs in large swarms L. pardalina scatters far and wide over the
central plateau of South Africa, but its natural home is in the semi-arid parts of the
country. It does not like the eastern Orange Free State, for instance, where there
is a fairly good rainfall and a rather dense growth of grass. Its favourable breeding
grounds are districts like those in the south-western corner of the Free State where
the rainfall is slight and the veld consists of short grass mixed with short Karroo
bush (Pentzia). In looking for scattered brown locusts I have got the habit of going
to spots in the veld where there is an outcrop of white limestone in the red sand.”

These valuable observations of Mr. Faure’s leave no doubt that L. pardalina
has, like L. migratoria, two different phases, which differ in morphology and colora-
tion, but more profoundly in the biology. Especially striking is it that there is a
sort of parallelism in the variation from the swarming to the solitary phase in both
these species, as will be presently evident. My study of extensive series of both
phases of pardalina sent by Mr. Faure, with a careful designation of the conditions
under which each particular lot was collected (i.e, whether from swarms or singly),
enables me to state the following differences between them.

The difference in the shape of the pronotum is well marked, though less striking
than that between migratoria (or mugratorioides) and danica. The pronotum of the
swarming phase of pardalina (fig. 5, A & B) is more constricted before the middle,
with the fore margin feebly prominent, the hind angle distinctly rounded, and the
median keel slightly lower, and deeper cut by transverse sulci than in the solitary
phase (fig. 5, D & E) ; but it is hardly possible to express these differences in figures,
as I have done for the phases of migratoria, in which they are far more pronounced.

It is possible, however, to apply the method of proportions to another character—
that of the relative lengths of the elytra and hind femora. As in migratoria, the
elytra of the swarming phase of fardalina are relatively longer and the hind femora

158 B. P. UVAROV.

shorter than in the solitary phase. The femoral proportion (7.e., the length of hind
femora expressed as a percentage of the length of the elytra) in the swarming phase
averages 0:44, with a maximum of 0-47, and a minimum of 0-41; the average
proportion for the solitary phase is 0:46, with 0-50 maximum and 0-41 minimum.

While the elytra of mzgratoria and danica do not differ except in the relative
length, there exists a well-marked difference in the shape and venation of the elytra
in the two phases of pardalina. Those of the swarming phase (fig. 5,C) are broader,
with the fore margin more convex and the apex obliquely rounded, while the solitary
phase (fig. 5, I*) has the elytra narrower, with the margins almost straight and parallel
and the apex obliquely truncate. The most striking difference, however, is in the
venation: the discoidal field in the swarming phase is much broader, and its false
vein distinctly curved and much thicker than in the solitary phase, in which this
field is rather narrow, parallel-sided, and with the false vein only slightly thickened
and almost, or even quite, straight. Naturally all these characters are subject to
variation, and forms in all respects intermediate occur,

ocseeaseeeacse
S-s52 ee
nizes, Sy

op. fobling

Fig. 5. Locustana pardalina, Walk.: A, B, C, phase pardalina ; D, E, F, phase solitaria, Uvar., nov.;
C and F, natural size, remainder x 3.

The difference in the absolute dimensions between the two phases of pardalina,
noted by Mr. Faure, is a phenomenon not observed in migratoria, but in the South
African species it is well marked, the average length of the body of the swarming
phase being 41 mm., while in the solitary phase it is only 34 mm., and occasional
specimens occur even as small as but 24 mm. The individual variability in size
is in the solitary phase very extensive, but the swarming phase is rather constant
in this respect; this is also the case in the phases of migratoria. Again, sexual
dimorphism is as well marked in the solitary phase of pardalina as it is in danica,
the average length of the body being 32 mm. in the males and 36 mm. in the females,
while the difference in the size of dwarf males, attaining about 22-24 mm., and the
largest females with a length of nearly 45 mm. is very striking. The males of the
swarming phase are scarcely smaller than the females, which agrees with the relative
size of the sexes in typical migratoria or migratorioides.

A REVISION OF THE GENUS LOCUSTA, L. 159

The extremely variable and decidedly protective general coloration of the solitary
phase of fardalina in all stages of post-embryonic development, noted by Mr.
Faure, exactly corresponds with the coloration of danica. The larvae of the swarming
phase of pardalina present an astonishing likeness to those of migratoria and migra-
totvoides, the coloration presenting the same combination of orange and black.*
The adults of the swarming form are rather uniformly coloured, and some bright
yellow specimens in the series sent from Pretoria lead me to the suggestion that
at the time of mating a general change of coloration may occur, as it does in migratoria
(see p. 143). In the coloration of the most aberrant individuals of the solitary
phase, one particular feature is noteworthy : the presence of a more or less pronounced
pale or white oblique cross on the pronotum, which design is very characteristic
of the species of the closely related genus Oedaleus.

An examination of the male genitalia revealed no difference whatever between
the two phases of pardalina, but this was only to be anticipated.

The above-quoted conclusions of Mr. J. C. Faure concerning the transformation
of the solitary phase into the swarming one, at which he arrived quite independently
of my work on migratoria, give a very strong support to the theory of phases as a
direct cause of the periodicity. His observations are especially interesting because
they concern the period of the transition from the solitary to the swarming phase,
on which my investigations of migratoria have given very few facts. It seems that in

De. pblieeg,

Fig. 6. Front view of head of: A, Locusta migratoria ph. danica, L.; B, Locustana pardalina
ph. pardalina, Walk. (x 4).

pardalina the transformation of solitary individuals into the swarming phase takes
more than one generation, but the actual causes of the transformation are in this
case also obscure, as they are in migratoria. Data as to the migrations of the flyers
and the fate of migrating swarms of fpardalina are yet lacking, and further
investigations of this problem, closely connected with the careful study of all
conditions of breeding grounds, are extremely important from the point of view
of locust control in South Africa.

V. SYSTEMATIC PART.

Key to the Genera Locusta, L., and Locustana, g. 1.

1 (2) Frontal ridge not widened at the median ocellus (fig. 6, A). Pronotum (fig. 1,
A, B, C, D, E) with the typical transverse furrow cutting the median keel
about its middle ; furrows in the prozona feeble. Mesosternal lobes only a

* It is extremely interesting to note here that the larvae of most swarming and migratory locusts
(Schistocerca peregrina, Ol., Dociostaurus maroccanus, Thb., etc.) present the same general type
of coloration in black and reddish, or yellow, forming a very striking design. This phenomenon
is well worth further investigation.

160 B. P. UVAROV.

little longer than broad. Elytra (fig. 1, F) not less than five to six times
as long as their maximal width ; hind radial vein diverging from the middle
radial only slightly and close to the bifurcation of the former ; discoidal
area much shorter than half the elytra ; inter-ulnar area about half as broad
again as the discoidal area, rather densely areolated, with areolets more
than three deep, without a regular false vein. Hind femora narrow, more
than four times as long as their maximal width ; their upper margin more
or less distinctly serrate ; upper carina of the externo-median area straight.
3: supra-anal plate (fig. 7, A) triangular, its surface practically flat ; cerci
(fig. 7, A) short, rounded, conical ; subgenital plate with apex obtusely
conical ; penis (fig. 7, B) very large, strongly recurved apically. § (fig. 7, E) :
subgenital plate with lateral margins straight ; apex truncate , lower |
valves of ovipositor with basal part distinctly longer than broad, with |
an obtuse lateral tooth in the apical part .. es ie Locusia, L. @
Genotype: Gryllus Locusta migratoria, L.

Fig. 7. Genitalia of Locusta migratoria, L.: A, male supra-anal plate and cerci from above ;
B, penis and lower genital valves (e) in profile ; C, epiphallus ; D, upper bridge of genital valves ;
FE, end of female abdomen from beneath. (E x 3, remainder x 12).

2 (1) Frontal ridge distinctly widened at the median ocellus (fig. 6, B). Pronotum
(fig. 5, A, B, D, E), with the typical transverse furrow cutting the median
keel distinctly before the middle; furrows in the prozona deep. Meso-
sternal lobes distinctly longer than broad. Elytra (fig. 5, C, F) not more
than four to five times as long as broad ; hind radial vein strongly diverging
from the middle radial long before its bifurcation ; discoidal area almost
as long as half the elytra ; inter-ulnar area about as broad as the discoidal,
sparsely areolated with two rows of areolets, separated by a false vein.
Hind femora broad, less than four times as long as their maximal width ;
their upper margin not serrate ; upper carina of the externo-median area
convex. dg : supra-anal plate (fig. 8, A) trapezoidal, with the apex prominent
in the middle, irregularly denticulate, its surface with chitinous tubercles ;

A REVISION OF THE GENUS LOCUSTA, L. 161

cerci (fig. 8, A) rather large, compressed laterally ; subgenital plate with
the apex slightly widened ; penis (fig. 8, B) very short, with short acute
apex. 9 (fig. 8, E): subgenital plate with lateral margins slightly convex ;
apex rounded, with a median projection and bisinuate laterally ; lower
valves of ovipositor with the basal part about as long as broad, the apical
part unarmed laterally oF 5 Locustana, g.n.

Genotype: Pachytylus pardalinus, Walk.

This key shows only the most striking differences between the two genera ; other
distinctive characters, especially those observed in the male genitalia, may be easily
understood by a comparison of the figures (figs. 7 and 8).

Fig. 8. Genitalia of Locustana pardalina, Walk.: A, male supra-anal plate and cerci from above ;
B, penis and lower genital valves (e) in profile ; C, epiphallus ; D, upper bridge of genital valves ;
E, end of female abdomen from beneath. (E x 3, remainder x 12)

A full description of the new genus Locustana is given below.

Key to the Phases of Locusta migratoria, L.* (figs. 1 & 4).

1 (2) Pronotum distinctly compressed laterally, but feebly constricted before the
middle ; median keel high, tectiform, convex in profile; the fore margin
angulately prominent; the hind angle acute. Hind femora (on the average)
longer than half the elytra. Hind tibiae often red. Larvae of variable
coloration, but never black and red (or black and yellow). Both larvae
and adults occur singly or in very loose swarms... ~-, ph. danica, L.

2 (1) Pronotum only feebly compressed laterally, but distinctly constricted before the
middle ; median keel low, neither tectiform, nor convex in profile ; fore
margin not prominent ; hind angle not acute. Hind femora (on the average)
shorter than half the elytra. Hind tibiae only exceptionally red. Larvae
of a constant black and red (or black and yellow) coloration. Both larvae
and adults occur in dense swarms.

* It is quite obvious that the exact determination of the phases is possible only in the case of
typical specimens and not of intermediate examples ; the most reliable results can be obtained
only from examination of large series.

162 B. P. UVAROV.

3 (4) Pronotum with the median keel straight in profile; hind margin rotundato-
angulate ie Bes ys A. as Py .. ph. migratoria, L.
4 (3) Pronotum with the median keel concave in profile; hind margin widely
rounded ey) he a hg: .. ph. migratorioides, Rch. & Frm.

Synonymic Notes.

The synonymy of L. migratoria ph. migratoria, L., and L. migratoria ph. danica, L.,
is quite correctly given by W. F. Kirby in his Catalogue (iti, pp. 229, 230), apart from
the fact that he distinguishes them as two different species, and I think it unnecessary
to repeat it here. To the synonyms of danica, however, must be added Pachytylus
australis, Sauss. (Prodr. Oedip., pp. 119, 120, no. 5, 1884), but not Locusta australis,
Froggatt (Agric. Gaz., N.S. Wales, xiv, p. 110, 1903), which represents Gastrimargus
musicus, F., as has already been stated by Prof. Y. Sjéstedt (Ark. Zool., xii, no. 20,
p. 11, 1920).

Pachytylus capito, Sauss. (Prodr. Oedip., pp. 119, 120, no. 4, 1884) is undoubtedly
identical with L. migratoria ph. migratorioides (Rch. & Frm.).

Pachytylus minor, Sauss. (Abh. Senck. Naturf. Ges., xxi, p. 631, 1899) belongs to the
genus Pternoscirta, as I am able to state from an examination of good photographs
of the type specimen most obligingly taken for me by Dr. J. Carl, of the Geneva
Museum.

The only remaining species of Locusta in Kirby’s Catalogue is L. pardalina
Walk., conspecific with sulcicollis, Stal, and capensis, Sauss., which is here made
the type of the new genus Locustana, m.

A Description of the Genus Locustana, nov. (figs. 5, 6 B & 8).

Antennae distinctly compressed dorso-ventrally. Frontal ridge in profile straight
or feebly concave, distinctly widened and impressed around the median ocellus,
flat elsewhere, with the margins very obtuse, disappearing just below the ocellus.
Fastigium of the vertex slightly sloping, forming a straight widely rounded angle
with the frontal ridge, flat, distinctly longer than broad, with the margins distinctly
raised, convex, with the median keel always developed ; the distance between the
eyes slightly less than twice as broad as the frontal ridge between the bases of the
antennae. Eyes oval, with the fore margin almost straight ; their width in the
broadest part, which is in the middle, is equal to about half their maximal height.
Pronotum with the prozona constricted, deeply furrowed, convex between the
furrows; the typical furrow cuts the median keel distinctly behind the middle ;
median keel moderately elevated ; lateral lobes with the hind angle widely rounded.
Mesosternal lobes distinctly broader than long, their inner angles widely rounded ;
mesosternal interspace subquadrate in the female and slightly longer than broad in
the male. Elytra hyaline throughout, except the basal parts of the marginal and
basal areas, which are coriaceous ; rather broad and short, not more than four to
five times as long as their maximal width ; apex oblique ; hind radial vein strongly
diverging from the middle radial long before its bifurcation (halfway between the
base and the bifurcation) ; discoidal area almost as long as half the elytra, with a
sinuate or straight false vein ; inter-ulnar area about as broad as the discoidal, or
scarcely broader, sparsely areolated, with two rows of the areolets separated by a
rather regular false vein ; axillar vein free, in most cases reaching the hind margin.
Wings rather short, not more than twice as long as their greatest width. Hind
femora rather broad—less than four times as long as their maximal width ; upper
keel not serrate; upper carina of the externo-median area distinctly convex.

$.—Supra-anal plate, with strongly chitinized margins, trapezoidal, distinctly
longer than the basal width ; its surface distinctly concave, with several small chitinous
tubercles in the basal half, forming a trapezium ; outer margins nearly straight ;

A REVISION OF THE GENUS LOCUSTA, L. 163

apex triangularly prominent, irregularly denticulate ; hind angles obtusely rounded.
Cerci rather large, strongly compressed laterally. Subgenital plate subconical, with
the apex attenuate and slightly widened. Penis* short, widely and obliquely truncate
posteriorly, with the apex triangular, sharp; upper genital valves large, united
with the penis ; lower valves small, lying close to the sides of the penis and covering
about half of it laterally, with obtuse upper projections ; they are connected with
each other above the penis by a bow-shaped transverse bridge emitting forwards
two long, apically narrowed, lateral branches ; epiphallus large, with the two upper
apophyses obtusely rounded and projecting inwardly and with lower sharply pointed,
beak-shaped teeth, with the inner lobes rounded and minutely and obtusely serrate
near the lower angles.

?.—Supra-anal plate obtusely triangular. Subgenital plate much longer than
broad, widened posteriorly, with the lateral margins slightly convex ; apex rounded,
bisinuate, with a small projection in the middle. Upper valves of the ovipositor
with short, strongly recurved, rather obtuse apices and not very sharp margins.
Lower valves with the basal part about as long as broad ; apical part without lateral
teeth, with widely rounded lateral angles; apices feebly decurved, short, acute.

Genotype: Pachylylus pardalinus, Walk.
Key to the Phases of Locustana pardalina, Walk.} (fig. 5).

1 (2) The average size smaller ; the males much smaller than the females. Pronotum
more compressed laterally between the shoulders, but less constricted
before the middle ; median keel more raised, not lower in prozona than
in metazona; fore margin more or less projecting in the middle; hind
angle sharp. Elytra shorter and narrower, with the margins almost straight
and parallel, with the apex obliquely truncate; discoidal area narrower,
parallel-sided, with the false vein straight and not incrassate. Coloration
of larvae, as well as that of the adults, very variable, but the larvae are
never coloured black and red. Both larvae and adults occur singly and
in loose swarms. ia = a " asd ph. solitaria, n. ph.

2(1) The average size larger; the males only a little smaller than females.
Pronotum not compressed laterally between the shoulders, but strongly
constricted before the middle ; median keel less raised, in prozona distinctly
lower than in the metazona; fore margin straight ; hind angle rounded.
Elytra broader and also longer, with the margins convex, and the apex
obliquely rounded ; discoidal area broader, with the sides sinuate, and the
false vein distinctly incrassate and sinuate. Coloration of the larvae
uniformly black and red. Both larvae and adults occur in dense swarms

ph. pardalina, Walk.

* The terminology of the parts of the genitalia adopted here is that of L. Chopard (Recherches
sur la conformation et le dévelopment des derniers segments abdominaux chez les Orthoptéres.—
Théses présentées a la Faculté de Sciences de Paris; Série A, No. 847, 1920).

t See the footnote on page 161.

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165

ON NEW SPECIES OF CURCULIONIDAE ATTACKING FOREST TREES

IN INDIA.

By Guy A. K. Marswattz, C.M.G., D.Sc.

Subfamily BRACHYDERINAE.

Sympiezomias beesoni, sp. nov. (fig. 1).

$2.—Black ; the head and rostrum with thin blue-grey scales; the prothorax
with rather sparse dull blue scales on the dorsum, the sides entirely clothed with
dense metallic green scales ; the disk of the elytra as far as stria 4 covered mainly
with blackish scaling, more or less interspersed with green scales, which are some-
times denser along the suture; beyond stria 4 the sides are clothed with dense
pale green scaling almost to the margin, the inner edge of the green area being very
irregular ; the lower surface with dense greyish green scaling.

Fig. 1. Sympiezomias beesoni, sp. n.

Head with coarse punctation that is usually confluent longitudinally, and often
with a very faint broad transverse impression behind the eyes; the vertex rarely
with a median stria, but usually with a central fovea. Rostrum a little longer than
its basal width, shallowlyimpressed in the middle, with the usual short median furrow,
which is not continued on to the forehead, the carinae on each side well marked and
parallel. Antennae piceous ; joint 1 of the funicle slightly longer than 2. Prothorax
a little broader than long, narrower in front than behind, broadest about the middle,
the sides gently rounded ; with coarse confluent punctation above, the interspaces

166 GUY A. K. MARSHALL.

finely and sparsely punctate, without any dorsal stria or impressions ; the setae
very short, flattened and recumbent. Elytra narrowly ovate m 4, only slightly
broader in 9, with the basal margin strongly raised and the humeral fold not very
prominent, the apices not mucronate in either sex; the striae strongly punctate,
the intervals slightly convex and of even height, except that interval 1 is somewhat
raised at the apex; the flattened setae short, recumbent and inconspicuous on the
basal half, longer and more curved behind. Legs with the hind tibiae not denticulate
internally.

Length, 7-8:75 mm.; breadth, 2-75-3-5 mm.

Mapras: S. Malabar, Nilambur, 5 ¢4, 14 99, vii.1918 (C. F. C. Beeson) ;
S. Malabar, Nadengayam, 2 $4, 1 9, xi.—xu1.1917 (N. C. Chatterjee).

Owing to its green scaling and non-mucronate elytra this species comes nearest
to S. frater, Mshl., which, however, has the whole upper surface with uniform green
scaling; the latter also differs in having no median impression on the rostrum,
the eyes are less convex, the pronotum is much more finely sculptured, and the
intervals on the elytra are flatter.

These weevils were found feeding on the leaves of young teak, and in some cases
defoliating the trees.

Subfamily ALCIDINAE.
Alcides dipterocarpi, sp. nov. (fig. 2).

$ 9.—Colour red-brown, the head, rostrum, prothorax and the humeral angles
of the elytra blackish ; the elytra very sparsely set with small narrow pale scales,
and without any squamose markings.

Fig, 2. Alcides dipterocarpi, sp. n.

Head coarsely and confluently punctate, with a shallow furrow just above each
eye, and the forehead broadly but shallowly impressed. Rostrum as long as (3)
or a little longer than (9) the front femur, parallel-sided from the base to the middle,
then narrowing slightly and widening again to the apex, which is a little broader
than the base; coarsely and closely punctate throughout, the punctures on the apical

NEW SPECIES OF CURCULIONIDAE. 167

half scarcely smaller than those at the base, with a shallow elongate median fovea
above the insertion of the antennae, and with a smooth median line on the apical
half only ; the sides clothed in the basal half with spatulate scales that are fringed
at the apex; ¢ without any projection on the submentum. Aztennae with joint 1
of the funicle about as long as the next three together, 4-6 subequal and as long as
broad, 7 densely squamose and a little longer than its apical width. Prothorax
nearly twice as broad as long, broadest at the base, the sides gently rounded and
markedly constricted near the apex, the dorsum closely set with low convex
granules, except the apical area, which is strongly punctate ; each granule bearing
a forwardly-directed seta (usually bifid) on its anterior edge, and the sides of the
apical area with numerous fan-shaped plumose scales, a few of which occur also on
the median basal lobe. Scwutellum small, transverse, tilted forwards, and not at all
enclosed in front by the elytra. Elytra broadly heart-shaped, scarcely longer than
the width at the shoulders, which are obtusely prominent ; the striae containing
round, deep punctures, which are for the most part separated by more than their
own width and become much shallower behind; the intervals broader than the
striae, convex, transversely rugose, but not granulate, and sparsely set with small
setiform scales, which are often bifid, trifid or quadrifid. Legs comparatively short
and stout, and thinly clothed with narrow, cleft scales ; the hind legs unusually
short, so that when outstretched the femur reaches only the apex of ventrite 6 (4th
visible), and the tibia exceeds the elytra only by its apical third ; the tooth on the
front femora of very unusual shape (fig. 3), being in the form of an irregularly trifid

Fig. 3. Front femur and tibia of Alcides dipterocarpi, sp. n.

lamina, that on the mid-femora truncate at its apex and with two minute teeth on its
external edge, that on the hind femora very small and simple ; the front tibiae with
a stout median tooth on the inner edge and a subapical one of almost equal size,
the mid-tibiae with a subapical tooth but no median one, and the hind tibiae simple.
Venter clothed with cleft scales, which only partly conceal the integument; ventrite 7
(last visible) with a large rounded median impression in the 3.

Length, 6-5-7-2 mm.; breadth, 4-25-4-8 mm.
UnitED Provinces: Dehra Dun, bred from seeds of Dipterocarpus, vii.1911.
Described from four specimens.

The rhomboidal outline and general structure make this species look like a very
small A. crassus, Pasc., from the Andamans; but the latter differs, inter alia, in
having a median internal tooth on all the tibiae, the intervals on the elytra are quite
smooth and sparsely punctate, the tooth on the fore femora is triangular with a
denticulate outer edge, and the ¢ has two tufts of setae on ventrite 7

There are several closely allied, and apparently undescribed, species in Malaya,

from all of which A. dipterocarpi may be distinguished by the peculiar form of the
tooth on the front femora (fig. 3).

168 GUY A. K. MARSHALL.

A. morio, Heller, from South India, is also of very similar appearance, but the
elytra are much less narrowed behind, the front coxae are closely approximate, and
all the tibiae lack the median tooth and the subapical one is inconspicuous.

Subfamily CRyPTORRHYNCHINAE.

Mecistocerus fumosus, sp. nov. (fig. 4).

gd 9—Colour black, with dense blackish or very dark sooty brown scaling
above, sometimes sparsely variegated with lighter brown scales ; the lower surface
with dense sandy brown scaling, the venter with a very broad, median, dark brown

stripe throughout.

Fig. 4. Mecistocerus fumosus, sp. n,

Head with the vertex bare and with coarsely reticulate punctures ; the anterior
part densely clothed with curled scales, completely hiding the integument, which is
much more finely sculptured than the vertex, bearing shallow larger punctures
interspersed with much smaller ones ; the frontal fovea very deep and elongate, the
scales between it and the eye mostly erect. Rostrum of 3 blackish in the basal half
and red-brown towards the apex, coarsely punctate towards the base and there
tricarinate dorsally, the apical part finely and sparsely punctate, the antennae
inserted at the middle; of 9, paler, a little longer and more slender, more finely
punctate throughout, and with the antennae inserted behind the middle. Antennae

NEW SPECIES OF CURCULIONIDAE. 169

red-brown, the funicle sparsely clothed with recumbent pubescence, the joints
clavate, the two basal ones equal, the remainder slightly and progressively diminishing
in length, all of them being longer than broad, except 7, which is about as broad as
long ; the club subcompressed, markedly broader than the funicle, as long as the
three preceding joints together, the first joint of the club as long as the remainder.
Prothorax with the sides subparallel from the base to the middle, thence roundly
narrowed to the apex and without any apical constriction ; the dorsum flattened,
with the usual very large and deep reticulate punctures, diminishing in size and
depth anteriorly, the interspaces shiny, not setose, and with squamiferous punctures,
without any real carina, but with a median undulating impunctate line that nearly
reaches the base and apex ; the scaling on the disk rather thin and recumbent, the
scales on the sides larger, denser and slightly raised, and a sharply defined line on the
pleura between the dark dorsal and pale ventral scaling. Scutellum trapezoidal,
broadest behind, about as long as its apical width, markedly flattened and sparsely
punctate. £/ytra unusually flattened, the shoulders prominent ; the posterior callus
distinct but obtuse ; the intervals plane, not elevated or granulate at the base, a
little broader than the large decp punctures, which are partly hidden by the dense
scaling ; when abraded the transverse septa between the punctures are seen to be
raised a little higher than the intervals between the rows: the scales very dense,
suberect, and with the tips curved downwards , the scale-like setae so short in the
male as to be hardly distinguishable from the scaling, a little longer and obliquely
raised in the Q. Legs with blackish scaling, except on the basal half of the femora,
where it is light brown ; the upper apical angle of the hind tibiae prominent, forming
a sharp right angle. Venter densely squamose at the sides, more thinly so in the
middle, with scattered raised squamiform setae ; ventrites 3, 4 and 7 (nominally 1,
2 and 5) with numerous large deep punctures, the interspaces with fine scattered
punctures , ventrites 5 and 6 each with a single transverse row of larger shallow
punctures.

Genitalia* of ¢ (fig. 5) with the median lobe almost parallel-sided, with a very

Fig. 5. Genitalia of Mecistocerus fumosus, Spum, Ga, aedoeagus ; 6, tegmen,

slight lateral sinuation near the apical third and the apex broadly rounded, only the
entire margin being heavily chitinised (probably immature) ; the uneverted sac

* The terms used are those suggested by Dr. D. Sharp, F.R.S. (Trans. Ent. Soc. Lond. 1918,
pp. 209-222.

(3442) N

170 GUY A. K. MARSHALL.

containing a pair of large lunate chitinous plates about the middle, the surface of the
sac around them being closely studded with asperities, and no plates at the median
orifice ; the tegmen with the ring not closed in the normal manner, but the chitinous
rim continued vaguely into the corresponding dorsal lobe on each side, the lobes
united for a short distance at the base and a little shorter than the length of the
ring, the strut being only half this length. Genitalia of 2 with a rather irregular,
oblong chitinous patch in the wall of the vagina opposite its junction with the oviduct,
and a similar broad V-shaped plate adjoining it anteriorly ; the spermatheca (fig. 7, d)

very broadly comma-shaped, the apex obtusely rounded, and the accessory gland
twice as long.

Length, 5-4-11 mm.; breadth, 2:2-4-8 mm.

UNITED PRovINcCES: W. Almora div., Kumaon, on Pinus longifolia (H. G.
Champion—type) ; Ranikhet, Kumaon, bred from P. longifolia (H. G. C.). PUNJAB:
Dalhousie Range, Chamba State, bred from P. longifolia (C. F. C. Beeson).

Its remarkable dark sooty colouring and flattened upper surface distinguish this
species from all its Indian congeners.

The species of this genus possess a stridulatory apparatus, which is the same in
the two sexes, and consists of a file formed of transverse striae at the apex of each
elytron on the lower surface ; the scrapers that play on these files are situated on
the seventh abdominal tergite and consist of two longitudinal rows of small, rather
widely spaced granules.

Genus Rhadinomerus, [st.*

{n his revision of the African species of Mecistocerus and Rhadinomerus (Ent.
Tidsk. xxv, 1904, p. 186) Prof. K. Heller treats the latter genus as merely a sub-
generic division of Mecistocerus. I agree with him that the supposed differences in
the structure of the venter used by Faust are quite unreliable; but this does not
apply to the form of the femora. In Mecistocerus these organs are clavate, being
markedly narrowed towards the base, whereas in Rhadinomerus they are sublinear,
being but little or not at all narrowed at the base ; moreover, as Faust indicated later
(Ann. Mus. Civ. Genova, xxiv, 1894, p. 279, note), the species of Mecistocerus can
be distinguished by the presence of an elongate bare shiny patch at the base of the
femora on the dorsal edge. But a more important point, which has hitherto been
overlooked, is that Rhadinomerus entirely lacks the apical stridulatory apparatus
that is present in both sexes of Mecistocerus ; and, finally, in females of the latter
genus the eighth abdominal tergite is as long as or longer than broad, whereas in
Rhadinomerus it is distinctly transverse. On these grounds it seems desirable that
Faust’s genus should be retained unless these characters are shown to be unstable.

Rhadinomerus bombacis, sp. nov. (fig. 6).

3¢.—Colour black; the prothorax with sparse, large, pale scales, which often form
a narrow median line and an indefinite lateral one on each side; the elytra with very
dense brown scaling, variegated with a few pale spots and transverse blackish

* Faust, Stett. Ent. Zeit. 1892, p. 215.

NEW SPECIES OF CURCULIONIDAE. 17]

patches ; the lower surface with a few sparse pale scales, especially towards the sides,
and obliquely raised flattened setae down the middle.

Fig. 6. Rhadinomerus bombacis, sp. n.

Head coarsely reticulate on the vertex, the forehead with dense recumbent scales,
which are larger behind than in front and partly conceal the median furrow, which
is rather shorter and shallower than usual. Rostrum with the usual four coarsely
punctate furrows and three narrow carinae at the base; the antennae inserted a
little in front of the middle in ¢ and a little behind it in 9. Antennae with the
funicle clothed with dense, erect pubescence ; the comparative lengths of the funicular
joints, beginning with the longest, are as follows: (1, 2) (3, 4) (5, 6) 7, all being longer
than broad except 7, which is as broad as long or slightly transverse ; the club only
slightly broader than the funicle, its first joint not longer than the second. Prothorax
a little broader than long, with the sides very gently rounded, broadest about the
middle, very slightly narrowed behind and much more so in front, with a shallow
constriction near the apex ; the dorsum, except the apical area, set with very large,
deep, reticulate punctures, and sometimes with a trace of a much abbreviated median
ridge ; the punctures on the pleurae smaller and much shallower and with the intervals
duller and more punctate; the pale scales much larger than those on the elytra,
oblong, convex and veined ; at the base of each puncture a suberect dark spatulate
seta directed forwards. Scutellum small, rounded, very convex and prominent.
Elytra subcordate, much broader than the prothorax at the shoulders, which are
roundly rectangular, with the sides subparallel from the shoulders to the middle
and rather abruptly narrowed behind, being markedly constricted at some distance
before the apex ; the large deep subquadrate punctures becoming much shallower
behind ; the slightly convex intervals as broad as the punctures anda little higher
than the septa between them, with indistinct granules, which are more evident near
the base, where the intervals are rather higher, each granule bearing a slightly raised
short scale-like seta. Legs with dense, light brown scaling; all the femora with a
faint, pale dorsal patch beyond the middle, behind which on the posterior pairs is
a large dark patch ; the tibiae with the basal half darker, except for a pale dorsal

(3442) N 2

72. GUY A. K. MARSHALL.

spot at the extreme base. Abdomen with coarse deep punctures on ventrites 3, 4
and 7 (nominally 1, 2 and 5), the punctures on the two former being denser on the
disk than at the sides; ventrites 5 and 6 impunctate and with the anterior edge
crenulated.

Genitalia of 3: the median lobe (fig. 10, c) oblong, troughlike and very slightly
narrowed towards the base, only lightly chitinised, except the inflexed lateral margins
and a transverse strip close to the apex, the latter area being dispersely punctate,
the apex broadly rounded, the base forming a sharp right angle on each side, and the
median orifice membranous and inconspicuous; the median struts slender, two-
thirds longer than the lobe, with a strong double sinuation in the basal third, and
connected together for a short distance at the base by an indefinite extension of the
lightly chitinised floor of the lobe ; the uneverted sac extending between the struts
to the end of the sinuated part, containing a small, median, lanceolate, chitinous
patch in the lobe close to its base, and the transfer-apparatus at the extremity of the
sac composed of two short juxtaposed hairpin-shaped pieces ; the tegmen (fig. 11, c)
with the strut a little shorter than the ring, and this again shorter than the lobes,
which are fused together for a very short distance at the base ; the spiculum twice as
thick as the median struts and slightly dilated at the apex, the basal fork forming
a very wide angle, one branch being twice as long as the other and abruptly curved
at its apical third. Genitalia of 2: spermatheca as shown (fig. 7, a).

Length, 3-8-6:5 mm.; breadth, 1-8-3-5 mm.

UniITED PRoviNcEs: Pathri, Sarahanpur, bred from logs of Bombax malabarica,
xi. 1917-1v.1918 (C.F. C. Beeson).

Brnar & Orissa: Singhbhum, from Bombax malabarica, 1.1921 (Beeson).

Described from 24 specimens.

‘Cc

Fig. 7. Spermatheca of (a) Rhadinomerus bombacis, sp.n.; (b) R. subfasciatus, sp. n. ;
(c) R. malloti, sp.n.; (d) Mecistocerus fumosus, sp. n.

Rhadinomerus diversipes, sp. nov. (fig. 8).

3 °.—Head and prothorax blackish brown ; elytra, venter and legs red-brown ;
the prothorax with the apical area densely clothed with pale brown scales and

NEW SPECIES OF CURCULIONIDAE. L/S

a few similar scales scattered over the disk, and with short, erect, dark setae;
the elytra rather thinly clothed with uniform pale brown scales (sometimes variegated
with paler spots), and with obliquely raised pale squamiform setae ; the sternum
with sparse, pale, fine erect setae, the venter with a very few pale scales and with
flattened, erect, pale setae.

Fig. 8. Rhadinomerus diversipes, sp. n.

Head bare and coarsely but shallowly reticulate on the vertex ; the forehead on
a slightly lower plane, with much shallower and smaller punctures, and with dense
pale scales; the frontal furrow very broad and deep. Rostrum tricarinate at the
base, as usual, with a small patch of raised pale scales in the middle of the base and
some erect, pale, squamiform setae; the antennal insertion at (9) or beyond (4)
the middle. Antennae testaceous, with the apical half of the funicle clothed with
sparse, recumbent pubescence ; the funicular joints in order of diminishing length :
(1, 2), 3, 4, 5, (6, 7) ; 5 as long as broad, 6 and 7 slightly transverse, the rest longer
than broad; the club with the basal joint a little shorter than the rest together.
Prothorax a little broader than Jong (11:9), with the sides rounded, broadest at the
middle, slightly narrowed to the base and shallowly constricted at the apex ; the
reticulate punctures not very large, but deep ; the interspaces dull, finely coriaceous,
and sloping inwards towards the puncture, each bearing a single erect, compressed,
dark seta; the median carina reduced to a very short sinuous line in the middle,
and sometimes almost obliterated. Scutellwm almost circular, strongly convex,
shiny, and with a few short recumbent hairs. Elytra subcylindrical, shallowly
constricted close to the apex and with an obtuse posterior callus ; the oblong punctures
deep and diminishing behind, each covered by an oblong horizontal scale attached
to the front margin, and on each side a small blackish prominence within the puncture ;
the intervals about as broad as the punctures, somewhat raised and rugulose close
to the base, but almost smooth elsewhere ; the scales varying from oval to oblong,
not sufficiently dense as to conceal the entire integument, and shorter than those on
the apical area of the prothorax ; the setae compressed, obliquely raised, and in
widely spaced rows. Legs with not very dense, uniformly pale brown scales ;_ the

174 GUY A. K. MARSHALL.

apex of the hind tibiae of the ¢ alone produced inwardly almost at right angles to
the tibia in the form of a lamina (fig. 9), having at its apex two short teeth representing

Fig. 9. Hind tibia of Rhadinomerus diversipes, sp. n. 3.

the uncus and mucro, the former being the longer. Abdomen with very large
separated punctures on ventrite 3 (Ist visible), the punctures being at most two
deep on the shortest portions (behind the coxae) ; ventrites 4-6 with only a few
much smaller punctures close to the lateral margins ; ventrite 7 coarsely and closely
punctate ; 5-7 with the anterior edge very broadly emarginate and bearing a close
fringe of short hairs, the emargination forming a well-marked obtuse angle at
each end.

Genitalia of 3: the median lobe (fig. 10, a) oblong, parallel-sided, and moderately

Fig. 10. Male genitalia (median lobe) of (a) Rhadinomerus diversipes, sp. n.; (b) R. malloti, sp. n.;
(c) FR. bombacis, sp. n.; (d) R. subfasciatus, sp. n.

chitinised throughout, except for an apical hyaline area which is broadly truncate,
the base sharply angulate on each side, the median orifice at about the middle of
the lobe, and the apical half with a few sparse punctures ; the median struts slender,
gently sinuous throughout, and half as long again as the lobe, with which they do not
definitely unite but merge gradually into the lateral edges of an abruptly narrowed,

NEW SPECIES OF CURCULIONIDAE. AS

indefinite, basal chitinous extension of the floor of the lobe: the uneverted sac
extending between the struts for three-fourths of their length, its terminal third
covered with asperities, the transfer-apparatus conspicuous and in the form of two
juxtaposed crook walking-sticks with the crooks turned outwards; the tegmen
(fig. 11, a) with the proportionate lengths of the strut, ring and dorsal lobes as
4:2-5:5, the lobes being fused together for one-third of their length from the
base ; the spiculum as broad as the tegminal strut, widely dilated at the apex, and
the basal fork forming a right angle with the branches nearly equal in length.

Length, 3-5-2 mm.; breadth, 1-4-2-4 mm.
UNITED ProvincEs: Lachiwala, Dehra Dun, bred from Eugenia yaman, x.1914
(C. F. C. Beeson—type}), and from Shorea robusta, xi.1915 (Beeson) ; Surajbagh,

Dehra Dun, bred from Eugenia jaman, xi.1915 (Beeson) ; Jubberkhet, Dehra Dun,
bred from Shorea, xii.1915-1.1916 (Beeson)

Described from 8 specimens.

Fig. 11. Male genitalia (tegmen) of (a) Rhadinomerus diversipes, sp.n.; (b) R. malloti, sp. n.;
(c) R. bombacis, sp.n.; (a) R. subfasciatus, sp. n.

Rhadinomerus malloti, sp. nov.

3§ °.—Head and prothorax blackish, the latter with yellowish brown scales on
the apical area only; the elytra piceous, irregularly mottled with lighter and darker
brown scaling; the lower surface without true scales, but sparsely set with short
and comparatively fine setae.

Extremely similar to R. diversipes, but distinctly broader in proportion to its
length. In that species the intervals between most of the punctures on the prothorax
are flattened and tilted inwards towards the puncture, so that the ridge is on the
outer or posterior edge ; in R. malloti these intervals are normally convex with the
ridge approximately in the middle. The scales in R. diversipes are oblong, with the
apex truncate or broadly rounded; in R. malloti they are narrowly ovate or lanceo-
late, with the apex pointed, and the setae are distinctly narrower. In the present
species ventrite 3 (Ist visible) is more closely punctate, the punctures being three
deep behind the coxa; ventrite 4 is also coarsely punctate, but less densely so than

176 GUY A K. MARSHALL.

3: 5and6 sometimes bear a single transverse row of punctures ; 5-7 have the anterior
margin only shallowly sinuate and without a fringe of hairs, the sinuation not being
angulated laterally. The hind femora bear a dark patch in the middle, the tibiae
are all darker on the basal half, and the hind pair in the ¢ have the uncus normal.

Genitalia of $: the median lobe (fig. 10, 6) with the sides straight and very
gradually diverging from the base to beyond the middle, then gently rounded to the
apex, which is broadly truncate and a little wider than the base, the lateral angles
of which are obtuse, the greatest width at one-fourth from the apex; the heavy
chitinisation is confined to the lateral margins and a slightly narrower transverse
band close to the apex, the rest being almost hyaline ; the median struts slender,
half as long again as the median lobe, gently sinuous throughout, distinctly
uniting with the sides of the lobe, and the connecting membrane between them
at the base hardly chitinised; the uneverted sac extending inwards for only
one-fourth the length of the struts and without apparent asperities, the transfer-
apparatus rather large and with two awl-shaped processes ; the tegmen (fig. 11, 6)
with the proportionate lengths of the strut, ring and dorsal lobes approximately
as 3:2:4, the lobes being united only at their extreme base ; the spiculum as wide
as the tegminal strut and only slightly widened at the apex, with one arm of the
basal fork about half as long as the other and forming an obtuse angle with it.
Genitalia of 9: the spermatheca as shown (fig. 7, c).

Length, 4-4-6 mm.; breadth, 2-2-75 mm.

Unitep Provinces: Lachiwala, Dehra Dun, bred from Mallotus philippinensts,
x.1914 (C. F. C. Beeson).

Described from 12 specimens.

Rhadinomerus subfasciatus, sp. nov.

g 9.—Colour piceous black, with the apex of the rostrum and tarsi paler; the
extreme base of the rostrum densely, the forehead less closely, clothed with cinnamon
scales ; the apical area of the prothorax with similar elongate scales, the remainder
not squamose, but with erect, compressed dark setae ; the elytra variegated with
more or less confluent spots of cinnamon-coloured scaling, which usually form a broad,
broken transverse band behind the middle, the dark areas thinly clothed with much
smaller dark scales; the lower surface without true scaling, but with sparse erect
pale setae.

The other external characters as described for R. diversipes, except the following :
Scutellum of similar shape, but quite devoid of hairs. Elytra with the setae a little
longer and distinctly narrower; the punctures not covered by a scale, this being
replaced by a very minute seta. Legs with the scales distinctly darker and smaller
on the basal than on the apical half of the tibiae; the hind femora with a dark
median patch; the uncus of the hind tibiae of the male normal. Abdomen with
ventrite 3 (Ist visible) much more closely punctate, the punctures being three deep
behind the coxae; ventrite 4 also bearing strong punctures, but much smaller and
more widely spaced than those on 3, and less numerous in the @ than in the @.

Genitalia of ¢: the median lobe (fig. 10, d) shaped just like that of R. mallott,
but the chitinised lateral areas more produced inwards in the middle, so that the
hyaline median area is shaped like an hour-glass ; the median struts slender, three-
fourths longer than the lobe, strongly bisinuous in the basal third, distinctly uniting
with the sides of the lobe, and connected together by a lightly chitinised membrane
in the basal fifth ; the uneverted sac extending inwards for nearly half the length
of the struts, set with asperities in the terminal area, but without any obvious
transfer-apparatus or other chitinous structure; the tegmen (fig. 11, d,) with the

NEW SPECIES OF CURCULIONIDAE. eA,

proportionate lengths of the strut, ring and dorsal lobes as 3-5 : 2: 3, the lobes fused
together for nearly half their length ; the spiculum similar to that of R. malloti,
but the fork forming almost a right angle. Genitalia of 9 : the spermatheca as shown
(fig. 7, 5).

Length, 3-5-4 mm. ; breadth, 1-5-2 mm.

UnitEp Provinces: Jhabberkhet, Dehra Dun, bred from log of Shorea robusta,
vi.l916 (C. F. C. Beeson—type); Kotdwara, Lansdowne Division, bred from
Shorea, 1x.1917. PunjaB: Thano, Siwalik Hills, bred from Eugenia, vi.1918
(Beeson).

The three previously described Indian species of Rhadinomerus (Faust, Ann.
Mus. Civ. Genova, xxxiv_ 1894 (1895), pp. 279-281) are known to me from description
only. Kk. granicollis is distinguished from all the species here described by its
granulato-punctate prothorax and the elytra bear spatulate suberect setae. R.
conciliatus is characterised by its cylindrical elytra, the intervals of which bear
minute granules and erect hairlike setae, the prothorax being oblong and parallel-
sided. The most distinctive features of R. contemptus appear to be that the prothorax
is roundly dilated before the middle and that it is densely squamose both at the sides
and at the apex.

Rhadinopus buteae, sp. nov. (fig. 12).

$ ¢—Colour black or red-brown, with dense, pale brown scaling and very broad
erect scale-like setae ; the pronotum with an indefinite blackish patch on each side
of the middle line near the base and a few white setae in the middle of the disk ;
the elytra each with an ill-defined oblique whitish band running from about the middle
of the suture towards the shoulder, but terminating on interval 5, and just behind
this on interval 2 an elongate patch of dense erect blackish setae ; the lower surface
rather thinly clothed with obliquely raised broad pale scales.

Fig. 12. Rhadinopus buteae, sp. n.

Head closely punctate on the vertex, each puncture containing a minute seta ;
the forehead not impressed, more coarsely punctate and with dense scales, which are
recumbent behind and erect in front, the interocular space parallel-sided. Rostrum

178 GUY A. K. MARSHALL,

rather strongly curved, strongly punctate at the base, and there rather more squamose
and with the median carina a little more prominent in the $; the antennal insertion
at (9) or a little beyond the middle (3). Antennae with joint 2 of the funicle a
little longer than 1, the remainder progressively diminishing and all longer than broad
except 7, which is as long as broad and bead-like ; the club very elongate, as long
as the 54 preceding joints, cigar-shaped. Prothorax broadest at the base, the sides
gently rounded, not constricted at the apex, the dorsum strongly and closely
punctate, without any median carina, the spaces between the punctures flat, shiny and
impunctate ; the scales large and subcircular, but scarcely overlapping, those in the
middle of the disk smaller and exposing more of the integument ; the very broad, erect
spatulate setae truncate at the apex. _ Scutellum almost circular, slightly convex and
densely clothed with small scales. Flytra ovate, broadest at the shoulders, which
form a rounded, obtuse angle, and scarcely impressed before the apex ; the compara-
tively small punctures almost hidden by the dense scaling, each one being covered by
a large scale and set in shallow striae ; the intervals a little broader than the striae,
not carinate on the disk, but rather convex and somewhat rugulose or subgranulate,
especially towards the base, only interval 9 carinate in the posterior half ; the scales
closely overlapping, and each interval with a row of very stout, erect, scale-like
setae, these being more numerous on intervals 2, 3 and 5. Legs with uniform pale
brown scaling and recumbent setae, the latter being erect only along the dorsal
edge of the tibiae; the dorsal edge of the femora straight and the inferior tooth
small; the tibiae markedly narrowed from base to apex, and with no angulation
externally at the apex.

Length, 6 mm.; breadth, 3 mm.

Punjab: Rani Range, Siwalik Hills, bred from logs of Butea frondosa, iv.1918
(C.F. C. Beeson).

Described from 8 specimens.

In the ¢ the last visible ventrite has a shallow impression at the apex, with a
small tubercle on each side of it bearing a single seta; in the § this ventrite bears
a shallow transverse impression at a little distance from the apex.

From the three previously described Indian species of the genus, centriniformis,
consputus and parcus (Faust, Ann. Mus. Civ. Genova, xxxiv, 1894 (1895), pp. 289-90)
the present species may be distinguished by the common pale V-shaped mark on
the elytra and by its tapering tibiae. R. centriniformis and consputus also differ,
inter alia, in having all the setae recumbent, interval 3 on the elytra strongly carinate
on the declivity, the dorsal edge of the posterior pairs of femora markedly sinuate,
the seventh ventrite (last visible) of the ¢ without tubercles, tergite 7 of the J almost
truncate at the apex and 8 densely squamose (in buteae 7 of the ¢ is deeply sinuate
at the apex and 8 is bare and shiny). R. parcus differs in having the pronotum
granulato-punctate, the scutellum is punctiform and shiny, the punctures on the
elytra bear only a fine seta instead of a broad scale, and the intervals are narrowly
carinate.

In this genus the stridulatory apparatus of the 3 consists of the usual files towards
the apex of the elytra near the suture, the scraper being formed by two very minute,
short transverse carinae on the apical edge of the seventh tergite, which are very
easily overlooked. In the 9 the files are on the seventh tergite, instead of on the elytra,
and are composed of comparatively widely separated longitudinal striae, the ridges
between them being very finely and transversely striate. The apparatus in both
sexes is similar to that foundin the European Cryptorrhynchus lapathi ; the statement
made by Dr. C. J. Gahan (Trans. Ent. Soc. Lond. 1900, p. 450) and repeated by
myself (Fn. Brit. India, Curculionidae, i, p. 17), that there are no stridulatory organs
in the 2 of this species is erroneous.

NEW SPECIES OF CURCULIONIDAE. 179

Subfamily ZYGOPINAE.
Osphilia odinae, sp. nov. (fig. 13).

3 2.—Colour piceous brown, rather thinly clothed above with pale hair-like scales,
asymmetrically variegated with subdenuded patches; the lower surface densely
covered with white scales, mostly linear, but some ovate; the base of the rostrum
and a broad stripe below each eye with dense, narrow, whitish scales. The pronotum
with pale yellowish scales, all of which are linear except a single row along the
extreme base, which are rather shorter and broader; a broad median denuded
stripe, and on each side of it a shorter oblong one narrowly uniting with it not far
from the apex, thus forming (very roughly) an inverted trident, externally to which
is a very indefinite lateral longitudinal sub-denuded patch; these darker areas
thinly clothed with dark recumbent setae. The elytra with the following similar
ill-defined and rather variable darker patches: a rounded one on the shoulder, a
transverse band before the middle extending from stria 2 nearly to the lateral margin,
a broad and very irregular macular transverse band behind the middle, and a rounded
juxta-apical spot.

Fig. 13. Osphilia odinae, sp. n.

Rostrum as long as the head and prothorax together, entirely red-brown,
gradually narrowed from the base to near the apex and thence slightly dilated, the
extreme base triangular in section with the apex uppermost; the basal portion
coarsely punctate (less so in 2), but the median line smooth throughout, the punctures
on the apical portion very fine and separated in 9, and longitudinally confluent in
3g; the antennal insertion behind the middle in 9, at or beyond it in g. Antennae
of 2 testaceous with only the club blackish, in the 3 the whole funicle also blackish
except joint 1 and the basal half of 2; the funicle with joint 2 as long as or a little
longer than 1, the remainder slightly and progressively shortening outwards in the 9,
all longer than broad, except 7, which is as broad as long ; in the ¢ joint 2 is strongly
clavate, the dilatation being greater on the inner side, which is densely clothed with
setae, the remaining joints bead-like and more nearly equal in length than in the @;
the first joint of the club longer than broad in both sexes. Pvothorax transverse,
the sides gently rounded, broadest before the middle, the base deeply bisinuate ;

180 GUY A. K. MARSHALL.

the dorsal outline almost flat, the dorsum finely but confluently granulato-punctate
throughout. Scwutellum circular, densely clothed with woolly scales having their
apices directed forwards. Elytra with the intervals broader than the very shallowly
punctate striae and finely rugulose ; the pale scales very long and narrow, except
on the basal half of interval 1, where they are much shorter and broader and pubescent
or densely fringed ; the scales on the darker areas short, setiform and dark. Legs
finely rugulose, rather thinly and uniformly clothed with long, narrow, pale scales ;
the front femora with a long sharp tooth, followed by a low carina bearing a single
row of 9 or 10 stiff erect bristles.

Length, 3-25-35 mm. ; breadth, 1-6-1:8 mm.

UnitepD Provinces: Banki, Gorakhpur, bred from Odina wodter, v.1918
(C. F. C. Beeson—type) ; Chauk, Gorakhpur, bred from Cassia fistula, v.1918
(Beeson) ; Dehra Dun, bred from Odina wodter, v.1919 (B. B. Osmaston).

Described from 17 specimens.

Only two other species of the genus have been described from the Indian sub-
region. O. brevirostris, Heller, from Ceylon, is of the same size, but has the rostrum
only as long as the prothorax, the joints of the funicle gradually widening outwards,
the antennal club pale and with the first joint broader than long, and the elytra clothed
with yellow scales and with a white spot at the apex of the suture.

O. egregia, Fst., from Burma, is a much larger insect (6 mm.) with mottled
umber and ochraceous scaling on the elytra, and three transverse rows of pale spots
on the pronotum.

181

INSECT PESTS OF VARIOUS MINOR CROPS AND FRUIT TREES IN
MAURITIUS.

By D. D’EMMEREZ DE CHARMOY,

Entomologist to the Department of Agriculture,

and S. GEBERT.

First Scientific Assistant, Division of Biology.

The following paper deals with insect pests of various minor crops and fruit
trees in the Colony of Mauritius. The damage caused by most of the pests dealt
with is often not very apparent, as many of these crops are not grown extensively,
and have not been the object of any special investigation up to the present. If
extensive cultivation of any of these should be undertaken, it is clear that there
might be created a new environment favourable to increase of the insects, and that
the pest problem would then have to be dealt with seriously. Our knowledge of
these pests is very deficient, as their life-history and habits have never been studied
in detail locally. It is also quite probable that extended cultivation may bring
to light many potential pests which at present escape our attention.

The crops dealt with in this paper are all capable of being developed in the future
and, for the present, nothing more than a mere enumeration of the pests for which
they serve as hosts can be made, except in a few particular cases. These brief notes
should only be considered as a preliminary study which will serve as a guide, as
well as a warning, to those who are likely to give more extensive attention to these
economic plants.

Cotton.

The cotton occurring in the Colony consists merely of a few plants scattered
over the island. In former years, as far back as 1847, the plant used to thrive in
many localities, where its cultivation was carried out on a moderate scale. The
yield and quality of the cotton were good, and no trouble due to insect attack seems
to have been experienced. Attempts to grow cotton on a large scale were made
in 1911, but owing to various unfavourable conditions, such as bad weather, drought,
scarcity of labour, and insect attacks, the enterprise did not succeed.

The Department of Agriculture, which did not exist then, is now taking up the
matter, and it is hoped that a certain extension will be given to cotton growing.

The enemies of cotton are mainly cosmopolitan, and most of the chief cotton
pests that have been recorded in other parts of the world occur here ; the following
have so far been noticed :—

Prodemia litura, F., and Spodoptera mauritia, Boisd. These two species of cut-
worms, though not true cotton pests, are nevertheless polyphagous, and therefore
likely to prove injurious to cotton. Pvrodenia litura is the commoner of the two,
and proves quite troublesome to garden plants during the hot season. The eggs
of this moth are parasitised by a species of Telenomus ; the development of the
parasite in the egg of the moth takes 21 days, and the proportion of males to females
emerging from the eggs is, on an average, 1 to 3.

Cosmophila flava, ¥. (xanthindyma, Boisd.). The caterpillar feeds on the leaves
of cotton.

182 D. D EMMEREZ DE CHARMOY AND S. GEBERT.

Chloridea obsoleta, F¥. This polyphagous insect of world-wide distribution,
commonly known as the American bollworm, is found here on a great variety of
plants, the chief being maize, tobacco, Hibiscus esculentus, Cajanus indicus, and

various other leguminous plants. It is also a serious pest of pelargoniums, destroying
the tender stems and flowers.

Earias insulana, Boisd. The caterpillar is the spotted bollworm of India. It
is another cosmopolitan species, and its feeding habits in Mauritius appear to be
much the same as those of Chloridea obsoleta. It seems, however, to confine its
attacks particularly to Malvaceae, and is of rather common occurrence. Twigs,
leaves and flower-buds are attacked as well as the bolls.

Platyedra gossypiella, Snd. The pink bollworm is one of the most serious pests
of cotton, and although it has not yet been observed on cotton in Mauritius, a few

moths have been bred from pods of Cajanus indicus and other Leguminosae of the
same type.

Pyroderces ( Anatrachyntis) simplex, Wlsm. The moth has been obtained from
larvae found in cotton bolls.

Dysdercus spp. Of the cotton pests which are found here, the red cotton-
stainers, of which two species exist, are by far the most common, and are found on
all malvaceous plants, as well as on many others, growing along the coast. As
cotton-stainers have done considerable damage in cotton-growing centres, they can
probably be looked upon as our most serious potential pest. Dysdercus is known
to transmit several bacterial diseases. It has already done a good deal of damage
to cotton bolls at the experimental plantation of the Department, and is the vector

of an internal disease of the bolls similar to that described by Nowell and others in
the West Indies.

Aphis gossypii, Glov. This green aphis is common on various garden plants
as well as on cotton. It is parasitised by a species of Ap/idius, and is preyed upon
chiefly by a large ladybird, Chilomenes lunata, and by Syrphid flies.

Satssetia migra, Nietn., and Sazssetia hemisphaerica, Targ. These two scale-
insects occur on various plants besides cotton. S. nigra is found chiefly on Hibiscus
esculentus, rubber, mango and various Malvaceae. It is parasitised by two species
of Chalcids and by a fungus. These probably keep it in check, as it cannot so far
be reckoned as a pest. S. hemisphaerica occurs chiefly on citrus, guava (Psidium),
litchi (Nepheliwm) and Aristolochia acuminata.

Achatina fulica. Though not an insect pest, mention may be made of this
large snail, locally known as “‘ Couroupa,” which also occurs in India and Ceylon,
whence it has been introduced here. It was responsible for a good deal of damage
to cotton seedlings, when the attempt was made to plant cotton on a large scale
in 1911. As the snails are very large, it is not difficult to have the land cleared of
them by hand-picking before planting. This should be done in the evening, as the
snail has nocturnal habits. Those met with in the daytime are generally found

hiding in dark places, or are in copulation. They are most active after the first rains
have set in and after sunset.

Tobacco.

Extensive cultivation of tobacco has recently been tried by this Department,
and as the results obtained have been very satisfactory, it is thought that tobacco
growing may rank among the chief of our minor industries. The variety tried is
that which is extensively grown in the island of Réunion. The yields: have been
satisfactory, and the prepared cigarette tobacco has sold rapidly. Planting is being
encouraged by the Department of Agriculture, which is doing everything it can in
order that tobacco cultivation may be carried out on a large scale.

INSECT PESTS IN MAURITIUS. 183

Although the insects attacking this plant are not numerous, it is nevertheless
useful to record them.

Prodenia litura, F., and Spodoptera mauritia, Boisd. The habits of these cater-
pillars have already been mentioned under cotton. The damage done to tobacco
seedlings may be very great.

Coelonia (Sphinx) solani, Boisd. A minor pest.

Phytometra (Plusia) orichalcea, ¥., and P. chalcytes, Esp. These two polyphagous
insects feed occasionally on tobacco. A certain amount of damage is caused to
seedlings.

Chloridea obsoleta, F. This moth is here the chief pest. The caterpillars eat into
the seed-capsules and often cause great damage ; they are also found on the leaves.

Thrips sp. Thrips are occasionally found in tobacco flowers, but do no great
harm.

Heterodera radicicola. The roots are sometimes so badly infested with eelworms
as to cause the death of a good many plants.

Maize.

Maize is another important food crop and its cultivation is being extended
yearly, there being now over 5,000 acres under this crop. Though the list of pests
is not long, appreciable damage is caused, especially to the cobs and seedlings.

Prodemia litura, F., and Spodoptera mauritia, Boisd. These two cutworms damage
the seedlings and are occasionally found on the leaves of the mature plant.

Sesamia vuteria, Stoll. Seedlings are badly attacked by the caterpillars of this
moth. Its life-history and habits have been worked out and published in a Bulletin
on Moth Borers of the Sugar-Cane (Scientific Series, Bulletin No. 5), issued by this
Department. Though Sesamia is a regular pest of sugar-cane, the moth has a marked
preference for maize seedlings, on which it deposits its eggs. As many as three or
four egg-batches are inserted between the leaf-sheaths and the stem; these are
composed of 15 to 75 eggs. The young larvae tunnel into the stem of the young
plant, which soon withers, and the caterpillars then migrate to older plants, in which

the rest of their development is passed, the caterpillar sometimes pupating in the
stem of the plant.

To gain an idea of the intensity of these attacks, the following is worth noting.
In March 1915, 35 acres of maize were planted on an estate, but so many of the
plants were destroyed that the yield amounted to that usually obtained from 11 acres.
On another occasion about 800 yards between lines of canes were planted with
maize ; the plants were examined every day, and showed the following infestation :—

Ist day of examination, 90 plants infested and uprooted.

2nd i, Hf 101 _ Me He
3rd i “ 88 i Ay -
4th ie » 64 x FA -
5th mi ie 164 ea . 4
6th Hy - 160 y ; ay
7th ee Pe 1s u ¥ ie
8th 2 ie 120 ‘, oy. .
9th A i 404 a at nd
10th rm - 1,925 ; Le a
11th + i 1,200 ™ v c
12th BS he 1,200 x i nA

Total 5,627 plants.

184 D. D’'EMMEREZ DE CHARMOY AND S. GEBERT.

It will be seen from this that Sesamia is a very serious pest. In addition to
boring the stems it also attacks the cobs. The natural enemies of this moth are
four in number. Telenomus sp. is an egg parasite, which has already been mentioned
under Prodenia. The life-cycle in the egg of Sesamia is the same as in that of Prodenia.
Trichogramma australicum, Gir., is another egg parasite. Henicospilus antancarus,

Morl., and Stauropodoctonus mauritit, Morl., are both parasitic upon the full-grown ,

caterpillar.

Proceras (Diatraea) sacchariphaga, Boisd., is less common than the former and
attacks the plant in the same way.

Aphis maidis, Fitch, is sometimes common on the leaves, but is not a regular
pest.

Chloridea obsoleta, F. The caterpillar is often found attacking the tender top
part of the stem and also the young ears. It frequently feeds on the silks, sometimes
destroying them entirely, in which case the ears become barren, and also bores its
way into the cob after having fed on the young grain. Many ears are spoiled in
this way.

When maize ears are allowed to remain on the plant until the latter has dried up,
the grain is invariably attacked by weevils, Calandra oryzae, L., being the chief pest.
Damage is also done by Dinoderus minutus, F. Over 50 per cent. of the grain may
be destroyed before storage. These pests, as well as a Tineid moth, also cause serious
damage to stored maize if fumigation is not resorted to. Full details concerning
these insects will be found in Bulletin No. 2 of this Department’s Scientific Series
(Insects Injurious to Stored Grains in Mauritius).

Manioe (Manihot utilissima).

Manioc is another important foodstuff, being largely used for stock. The plant
thrives in the hot districts, and gives very good returns. A great many varieties,
most of them imported lately by the Department of Agriculture, are grown. There
are practically no insect pests of the plant.

Lachnosterna (Phytalus) smith, Arrow, is an occasional pest in the infested
regions.

Saissetia hemisphaerica, Targ., is sometimes found on the leaves, but is a minor
pest. A species of Chionaspis occurs frequently on the stem.

Sweet Potato (/pomoea batatas).

This tuber is greatly in favour with the poorer classes of the population, and is
also an important cattle food. The plant grows well all over the island, the highest
yields being obtained in the hotter lowland regions. A great number of varieties
are grown, most of which have been imported lately by the Department of Agriculture.

The chief pest of the foliage is a Pterophorid moth, Trichoptilus wahlbersi, Z.,
the larva of which feeds on the leaves, especially the tender parts and buds, and
rolls up in the leaf before pupating.

Ercta ornatalis, Dup. This Pyralid moth is another leaf pest.

Herse (Sphinx) convolvuli, L. The caterpillar is occasionally found on the sweet
potato, but cannot be regarded as a pest.

Aspidomorpha obovata, Klug. This Chrysomelid beetle is also an occasional pest,
doing no great damage. It lives on various wild species of Jpomoea.

Cylas formicarius, F., is by far the worst pest. It is exceedingly common whenever
sweet potatoes are grown in mounds, and is then responsible for very serious damage.
As many as 50 to 100 larvae may be found in a single tuber. The method of planting
sweet potatoes in mounds has only recently been adopted, and the resulting looseness
of the soil certainly renders the plants more liable to the attacks of this beetle.

INSECT PESTS IN MAURITIUS. 185

Ambrevades (Cajanus indicus).

This plant suffers a great deal from caterpillars, which live in the pods. In
certain regions 75 per cent. of the crop is lost. Control measures are difficult to
apply, for reasons mentioned further on.

Adoretus versutus, Har. This beetle, which has polyphagous habits, feeds on
the leaves of the pigeon-pea, but is not a serious pest.

Lachnosterna (Phytalus) smith, Arrow. Plants grown in the infested region are
usually visited by L. smitht. The damage caused is insignificant, as the plants in
the infested areas serve as trap plants, from which the beetles are hand-picked at
night.

Icerya sevchellarum, Westw., is common enough on almost all the plants of the
island, and is only a minor pest of pigeon-pea. Various insecticides in the form
of emulsions have been tried against this pest, but the results have never been
successful.

Eucalymnatus tessellatus, Sign., is a minor pest of the twigs and attacks various
other Leguminosae.

Chionaspis subcorticalis, Green, is occasionally found on the branches of this
plant, more especially on the tender twigs, but does not do much harm.

A species of Botys is very common on the tender leaves.

By far the worst pests of Cajanus indicus are certain Lycaenid butterflies, which
are exceedingly common when the plant begins to flower. +The eggs are laid on the
flower-buds and young pods, and the newly-hatched caterpillar, after having fed
for a short time on the surface of the buds and pods, bores its way in. Advantage
was taken of this to try to control the insect by spraying the flower-buds and
pods with arsenate of lead. The results were not successful, for it was difficult to
make the spray adhere to the flower-buds, as these are covered with fine hairs, which
will not hold the spray. Moreover, at this time of the year the wind is usually very
strong, so that the sprayed material is shaken off the plant before it has time to dry,
in spite of its adhering power having been increased by the addition of mucilaginous
substances, such as a maceration of Opuntia in water. The following is a list of
the Lycaenids that feed in the flower-buds and pods :—Lampides boetica, L., Zizera
lysimon, Hb., Tarucus telicanus, Lang, and Nacaduba manderst, H.H.D. Of these
Lampides boetica is the worst pod pest, the others confining their attacks rather to
the flower-buds.

Chloridea chsoleta, F., is an occasional pest of pods.

Platyedva gossypiella, Snd., is also occasionally found in pods and is more common
than Chloridea.

Botys spp. Two species of PyRALIDAE are found in the flower-buds, and more
rarely in the pods. These are the most dangerous pests of flower-buds, and are
responsible for a great deal of damage.

Pois Sabre (Canavalia ensiformis).

Only a few pests are worth recording :—
Thrips are often abundant in the flowers and cause many of these to fall off.

Argyroploce rhynchias, Meyr. The caterpillar feeds in the pods, also attacking
the twigs, and may sometimes ruin the whole crop.

Aspidiotus sp. This scale is a minor pest, and is found on the twigs.

Rhopalocampta forestan, Cram., is also a minor leaf pest.

(3442) o

186 D, D EMMEREZ DE CHARMOY AND S. GEBERT.

Voheme (Vigna catjang) and Haricot Bean (Phaseolus vulgare).

Agromyza phaseoli, Coq., is such a regular pest of these plants that their
cultivation is almost impossible in certain localities and at certain times of the year.
Remigia (Pelamia) repanda, F¥. The caterpillar feeds on the leaves of Vigna
catjang.
Pea (Pisum sativum).

Plusia orichalcea, ¥. The caterpillar is a minor pest of peas, doing some injury
to the leaves.

Chloridea obsoleta, F., occurs as a leaf pest.

All the Lycaenid butterflies mentioned under Cajanus indicus attack the pea
more or less. The chief pest, however, is Lampides boetica, L., which is sometimes
responsible for great damage.

Pistache (Avachis hypogaea).

The pests attacking ground-nuts are not numerous and do not cause very serious
damage.

Chloridea obsoleta, F., and Plusia orichalcea, F., are the only two worth recording
as leaf pests. Ephestia cautella, Walk., attacks the fruits, and Pseudococcus
calceolariae, Mask., is sometimes a serious pest of the roots.

Pumpkins, Melons, Cucumbers, etc.

Cucurbitaceous plants suffer severely from the attacks of the fruit-flies, Dacus
sygmoides, Coqg., and Tridacus d’emmerezt, Bezzi.

The caterpillar of a Pyralid moth, Glyphodes indica, Snd., feeds on the tender
leaves and buds. It is not greatly harmful to the plant itself, but as the leaves are
used for human consumption, and as the caterpillars always roll themselves in the
leaves, they are troublesome pests.

Aphis gossypi1, Glov., is a minor pest.

Cruciferae (Cabbage, Cauliflower, etc.).

The Pyralid moth, Crocidolomia binotalis, Z., has proved to be a regular pest
of cabbage. The leaves are so badly eaten that they rot i situ and are thus rendered
unfit for human consumption.

Plutella maculipennis, Curt. Late in the season the leaves of cauliflowers and
cabbages are badly attacked by this pest.

Aphis brassicae, L., occurs in abundance at the beginning of the season. Towards
the end its numbers are reduced, as it is preyed upon by Syrphid flies, chiefly
Xanthogramma pfeifferi, larvae of lace-wing flies, and by a large ladybird, Chilomenes
lunata.

Artichoke.

Two pests occur :—The artichoke moth, Porpe bjerkandrella, Thunb., which is
very prevalent at times and greatly injures the foliage ; and an Aphid, Macrosiphum
picridis, F., which is exceedingly common on the leaves and tender parts of the
stem, doing serious damage.

Tomato.

The insect enemies of the tomato are the same as those of tobacco and need
not be mentioned again. It is worth recording, however, that the eel-worm,
eterodera radicicola, is a regular pest and sometimes occasions great losses.

INSECT PESTS IN MAURITIUS. 187

Coconut.

This plant thrives in the coastal regions wherever the land is not rocky. A certain
amount of extension is being given to its cultivation in certain parts of the coastal
belt, and promising results are being obtained.

The beetle, Oryctes tarandus, Oliv., occasionally bores into the tender part of the
stem, which it usually enters beneath the basal part of a leaf, just where it begins to
stretch away from the stem, and bores a gallery right through. It also bores its
way into the mid-rib of the leaf. The galleries in the stem are usually made
tangentially, and in such cases, though the tree is not immediately affected thereby,
a door is opened for bud-rot. It often happens, however, that the beetle bores its
way radially inwards into the growing point ; in such a case the health of the palm
is affected and immediate death results.

Of all the scales which are found on the coconut palm, the most common is
Diaspis boisduvali, Sign. It occurs in very great numbers all over the island and
is a serious pest. The injury done is very great both to old and young plants, the
insect occurring in thousands on a single leaf. Now that a certain extension is being
given to the cultivation of the coconut, combative measures will have to be adopted,
as the pest is a serious menace to the future of the coconut industry.

Coffee.

Coffee was extensively grown years ago, but its cultivation has had to be aban-
doned on account of the attacks of leaf disease (Hemileia vastatrix). It is, however,
cultivated in certain localities for local consumption. Liberian coffee thrives in
the colder districts and yields heavily, and if its cultivation is extended good results
may be expected, as it is very slightly subject to attacks of Hemileia.

Cratopus punctum, Boh. Arabian coffee does not suffer much from the attacks
of this weevil, which has a marked preference for the Liberian variety, the leaves
of which it devours to such an extent as to cause almost entire defoliation of the
plant. It is worth noting that in the upland regions the attacks of Cvatopus are
not serious.

Adoretus versutus, Har., is another leaf pest.

Botys octoguttatus.* The caterpillar feeds in the berry and is rather a serious
pest.

Prodemia litura, ¥., destroys many seedling plants as they come up.

Saissetia nigra, Nietn., and Saissetia hemisphaerica, Targ., occur as minor pests.

Coccus viridis, Green, is sometimes found abundantly on the leaves of young
plants in nurseries. Infestation begins as soon as the first leaves are fully developed
and soon extends to the young leaves as these emerge from the bud stage. In such
cases the young plants soon die. The insect also occurs in abundance on older
plants, being the most important scale pest.

A small Bostrychid beetle tunnels into the twigs of the coffee plant. The galleries
are bored obliquely to a length of 2 to 3 cm., reaching the pith. The attacks are
sometimes so bad as to cause the drying up of the infested twigs.

Limes and other Citrus Fruits.

The lime (Citrus medica var. acida) thrives in many parts of the island, especially
in the hotter coastal regions. The crop forms the subject of a small trade on the
local market, but is not of sufficient size to be dealt with on an export basis. Experi-
mental trials are being made at present by the Department of Agriculture with a
view to extending the cultivation of this plant, and it is hoped that before long lime

* (Possibly intended for Thliptoceras octoguttalis, Feld.—ED.]
(3442)

188 D. D’EMMEREZ DE CHARMOY AND S. GEBERT.

products will rank among the chief of our minor industries. Preserved limes are
imported from Rodrigues, a dependency of Mauritius, where the trees bear good
crops. The crop obtained there is, in fact, too large for the wants of the island,
and the excess is salted and sent to Mauritius. Other citrus fruits, such as mandarines,
oranges, lemon, shaddock, etc., also grow well and give good returns, so that on the
whole the group is of increasing importance locally.

There are many insects that attack these plants, the chief pests being Papilio
demodocus and the citrus aphis.

Papilio demodocus, Esp. This butterfly, which is known all over Africa as an
enemy of citrus trees, is here a most troublesome pest to young plants. The cater-
pillar destroys the seedlings when they are three or four inches high, and if removal
of the larvae from young plants is not practised constantly, considerable damage
is done. No great injury is done to full-grown plants. The eggs and larvae are very
conspicuous and can be easily removed from seedlings by hand-picking.

Papilio phorbanta, L. The life-history and habits of this butterfly are much
the same as those of Papilio demodocus. It is less common than the latter and
can rarely be reckoned as a serious pest of seedlings. The eggs are always laid on
the lower surface of tender leaves; Papilio demodocus, on the other hand, seems
to deposit its eggs anywhere.

The scale-insects found on lime trees are eleven in number and occur in abundance
during the beginning of summer. When the rainy season sets in and the air becomes
saturated with moisture, the scales are to a great extent attacked by several parasitic
fungi, which gradually destroy them. This statement does not, however, always
hold good for the coastal regions, where the heat is excessive and the climate com-
paratively dry. It is especially there that the presence of scale-insects is felt, for
the damage caused is appreciable and the scales occur all the year round. The ten
species found are :—

Chrysomphalus aurantit, Mask., and Chrysomphalus ficus, Ashm., on the twigs
and stems.

Pseudaonidia trilobitiformis, Green, on the leaves.

Chionaspis citri, Comst., on the leaves and tender branches.

Lepidosaphes gloveri, Pack., on the leaves, tender shoots, and fruits.

Saissetia oleae, Bern., on the young shoots and branches.

Saissetia hemisphaerica, Targ., on the leaves.

Coccus viridis, Green, on the leaves and tender twigs. Infestation is sometimes
very heavy, two to three hundred larvae and adults occurring on the same leaf.
This species is heavily parasitised by two Chalcids, Diversinervus silvestri1, Waterst.
sp. n., and Tetrastichus sicarvus, Silv.

Icerya seychellarum, Westw., is a regular pest.

Pseudococcus citrt, Risso, is a minor pest.

Pseudococcus filamentosus, Ckll. (vastator, Mask.) is commoner than the preceding
species.

The buds and flowers are intensely attacked by a species of brownish-black aphis.
The damage caused to the flowers is great, as a large proportion of them drop. Seed-
lings are sometimes infested to such an extent as to cause their growth to be stunted
for a time. When such is the case, lateral growths are emitted, to the detriment
of the plant.

Eggs and larvae of a Psyllid, Trvioza sp., are found on the lower surface of the
leaves at the beginning of summer. The larvae are greenish yellow and reach the
adult stage in about a month. Wherever the larvae occur there is a corresponding
swelling of the tissue, resembling a gall, on the upper surface of the leaf. The body
of the larva fits exactly into this pseudo-gall, its dorsal part lying flush with the
lower surface of the leaf. Tvioza is not a serious local pest, though it occurs sometimes
in appreciable numbers. When such is the case, the attacked leaves fall off, though
not until the insects have reached the adult stage.

INSECT PESTS IN MAURITIUS. 189

Mandarines suffer every year from attacks of the fruit-fly, Ceratitis catotrt, Guér.
The only remedial measure against this pest would appear to lie in the introduction
of parasites.

Mango.

Batocera rubus, L. The larva of this Longicorn beetle causes appreciable damage
to various trees, amongst which is the mango. In certain cases trees have been
found to suffer so severely that control measures had to be adopted.

Eight to ten years ago a Cecidomyiid fly, Procontarinia matteiana, Kieff., found
its way into the Colony. It is known to occur also in India, and was probably intro-
duced accidentally from that country. The insect is extremely destructive, and
wherever it is found the yield is reduced enormously, the leaves being infested when
they are still quite tender and about 2 inches long (fig. 1). At this stage small spots

cow Th eS Ses en A ee ee

!
nant Pi crt aes hg a a al ea

Fig. 1. Lower surface of mango leaf showing galls made by Procontarinia mattetana, Wieft.,

denote the places where the insect has oviposited. As the larva grows up in the
leaf tissue, a gall is formed ; the one harbouring the adult insect prior to its emergence
measures from 1-5 to 2-0 mm. in diameter. The gall shows equally on either surface
of the leaf, but the fly emerges by puncturing the lower surface. Infestation is
usually very severe, as many as 150 to 300 galls being formed on a single leaf. They
occur in such large numbers that they touch one another, there being scarcely room
for more. The leaves affected in this way soon dry and fall off. The life-cycle of
the insect is about two months, its period of most common occurrence being from
November to June inclusive.

Coccus mangiferae, Green, is quite common and sometimes occurs in great
numbers. After the Cecidomyiid, it certainly is the worst pest of the mango leaf.
The other undermentioned CoccrpaE cannot as a rule be regarded as regular pests.
These are :—Coccus hesperidum, L., Eucalymnatus tessellatus, Sign., Chionaspis
dilatata, Green, Pseudaonidia trilobitiformis, Green, Vinsonta stellifera, Westw., and
Icerya seychellarum, Westw. The last-named is sometimes quite common and does
a certain amount of injury. As a result of the combined attacks of scale-insects,
the leaves become covered with their honey secretions. On this covering there is
a dense growth of sooty mould (Capnodium sp.), which obstructs the stomata. The
leaves have a miserable appearance and cannot carry on their physiological functions
as they should, so that the whole tree suffers.

190 D. D' EMMEREZ DE CHARMOY AND S. GEBERT.

The larva of the mango weevil, Cryptorrhynchus mangiferae, F., feeds in the ~

stone, where it pupates. It is not a seriously destructive pest.

Ceratitis catoiri, Guér. Acertainamount of damage is caused by the larva
of this fruit-fly.

Peaches.

Adoretus versutus is more or less common all over the island, and frequently
attacks the leaves.

Aulacaspis pentagona, Targ., occurs on the stems and twigs. In the hot coastal
regions this pest is the worst of those affecting this plant. It is extremely common
and causes the death of a great number of trees every year.

Cydia pomonella, L., is another regular pest and is most harmful to peaches.
On an average 75 per cent. of the fruits are infested, especially at the beginning of
the ripening season.

Banana.

Bananas are grown all over the island, and many varieties are cultivated, amongst
them being the “Nain” and “Ollier.” The “ Gingell ” is affected with a
disease of the fruit in the colder districts, but the other varieties thrive all over
the country and form one of the chief food-stuffs of the poorer classes. They con-
stitute the most popular fruit of the island, as they are cheap and procurable during
most of the year.

The black banana weevil, Cosmopolites sordidus, Germ., is the only pest. The
larvae live in the root-stock and occur sometimes in such great numbers as to cause
the death of the plant. The varieties known as “ Gingeli’”’ and ‘‘ Banane Carrée 4
are particularly affected.

191

THE EGG PARASITES OF THE COFFEE BUG (ANTESTIA
LINEATICOLLIS, STAL) IN “KENYA COLONY;

By F. W. Dry, MSc.,

Recently Assistant Government Entomologist, Kenya Colony.

I. INTRODUCTION.

Some account of work on the egg-parasites of the coffee bug in Kenya Colony
has already been published in a bulletin of the Department of Agriculture of that
country.*

When that bulletin was written the two common species of egg-parasites had
not been identified, and they were consequently referred to simply as ‘ Species A,
Brown Parasite,’ and “‘ Species B, Black Parasite.’’ These two species both proved
to be new and have since been described by Dodd,f ‘‘ A” as Hadronotus antestiae,
and “B” as Telenomus truncativentris. They may be roughly distinguished as
follows :—

Hadronotus. Abdomen brown, head and thorax black in the female; head,
thorax, and abdomen black in the male; black colour dull; antennae in both
sexes of 13 segments.

Telenomus. Head, thorax, and abdomen bright glossy black in both sexes ;
abdomen of a different type from that of Hadronotus ; antenna of the female
12-segmented and comparatively short and clubbed, antenna of the male
13-segmented, comparatively long and moniliform.

It is now possible to give some additional facts, but it should be made clear that
the investigation is in a very incomplete state. This account, put together upon the
present writer’s ceasing to work in Kenya Colony, is a short summary of what has
so far been ascertained.

II. THE LIFE-HISTORY OF THE COFFEE Buc.

A full account of the life-history of the coffee bug is given by Mr. Anderson.
The following facts, the chief ones about the life-cycle that have a bearing on the
relation between the bug and its egg-parasites, are taken from his bulletin.

The egg stage and all the instars of the nymphal stage were passed through
considerably faster in hot than in cool weather. The length of the egg stage varied
from about 9 days in the hotter months to about 13 days in the cooler ones. The
average length of the nymphal stage was, for the hotter months, about 75 days,
for the colder, 115 days. Combining egg and nymphal periods, the average times
from egg-laying to the nymph becoming adult were respectively 84 days and 128 days.
The average adult life for both sexes was a little over 100 days, but the length of
life was frequently much longer, the maximum found for a female being 290 days,
that for a male 249 days. The average number of eggs laid during the lifetime of
a female was 126, but the number was often very much more, 485 being the highest
obtained.

The egg of the coffee bug, as Mr. Anderson describes it, “is dull white in colour,
covered with a fine powder which easily rubs off, leaving the egg clear and glistening.
This powder appears in the form of a delicate reticulation. The egg measures
roughly 3 mm. by 1 mm. The measurement of 100 eggs, taken at random, gave

* Anderson, T. J.—The Coffee Bug.—B.E.A. Dept. of Agric., Div. of Ent., Bull. i, 1919.
g Pp §

+ Dodd, A. P. Notes on the Exotic Proctotrupoidea in the British and Oxford University
Museums.—Trans. Ent. Soc. Lond., Jan. 1920.

192 F. W. DRY.

the following averages :—shorter axis 0:87 mm., longer axis 1-125 mm., the longer
axis being at right angles to the point of attachment. As the embryo develops
within the egg a clear ring becomes distinct on the top, marking the place where
the cap will split off when the young nymph emerges. The cap is a complete arc
of the egg-shell and may be completely split off or left behind, hinged on, and
may shut down accurately. In some instances it fits down so tightly that only the
colour shows whether the nymphs have hatched. When the egg is from four days
to a week old, depending on the season, the eyes and tylus of the nymph can be
distinguished as darker spots shining through the cap. The eggs are usually found
in clumps of twelve, though ten and eleven are quite common. The clumps of eggs
are found on the under surfaces of the leaves, occasionally on the upper surface,
on the berries, on the pedicels of the berries, on the stem, on dry leaves, and even
on stones beneath the bushes.”’ In the field 90 per cent. of the eggs collected were
found on the lower side of the leaves.

Ill. PARASITISED: EGGS.

The female parasites have been watched vigorously prodding the coffee bug eggs
with their ovipositors. The insect remains quiet for some time with the ovipositor
embedded in the egg, the whole operation lasting from two to five minutes. Eggs
that have been parasitised are distinguished by the fact that some time after being
attacked by the parasite, this time varying with the season, the colour of the egg
changes from the original dull white. Usually the eggs become bluish grey, but
occasionally parasites emerge from eggs which have only become pale grey.

The parasite makes its way out of the egg through a hole with a ragged edge
bitten by the insect’s mandibles; the cap of a parasitised egg does not split off.
Many hundreds of eggs have been kept singly in tubes in the laboratory, but
never more than one parasite has been reared from one egg.

In addition to the two named species of parasites other species have been reared,
but their numbers have been very few.

In collections brought in from the field the eggs were kept under observation
so as to determine how many produced ntestia, how many had been parasitised,
and how many failed both to turn blue and to produce either bug or parasite. All
eggs which turned blue were recorded as parasitised, and were then kept in tubes
plugged with pads of cotton wool. The parasites hatching from them were preserved
and examined, and data for proportions of species and sexes are given below. Some
of the eggs which turned blue did not produce parasites. Any eggs about which
there was a doubt were kept until they produced parasites or until it was clear
that they were not going to produce anything.

In making counts it was the practice to reject all eggs which had hatched—
whether Axtestia or parasites—before being collected, and to reject as well all clumps
containing one or more eggs which had hatched before coming to hand. In this
way, for each collection of eggs examined, percentages were obtained of those hatching
Antestia, those parasitised, and those not going blue or hatching anything. The
figures so calculated may be taken to represent fairly well the parasite position
in the plantations where the collections were made, but the following sources of
inaccuracy must be noted :—-

(a) A parasitised egg remains unhatched longer than one which produces a coffee
bug.
(b) If the eggs had not been brought into the laboratory some which hatched

Antestia would have been parasitised in the field; this tends to counter-
balance (a).

THE EGG PARASITES OF THE COFFEE BUG IN KENYA COLONY. 193

In discussing the data from collections of eggs we may conveniently divide

these collections into two lots :—

A. Those from a field of coffee about six acres in extent on the Government Farm
at Kabete. From this field a considerable number of collections have been
made over a period of two and a half years. Particulars from these collections
will be given first.

B. Those from various other coffee plantations in the districts of Nairobi, Kyambu,
Limuru and Thika, but on these plantations usually only a single collection
has been made, and that generally at the time of an Antestia outbreak. It
will not be necessary to say much about the data from these occasional
collections.

IV. COLLECTIONS FROM GOVERNMENT FARM, KABETE.

Between July 1917 and September 1920 twenty-four collections were made
from this field in as many different months. The eggs collected totalled 16,531, and
the months in which they were obtained are shown in Table I.

The chief points brought to light are the following :—

The average of the monthly percentages of eggs producing Antestia was 15 per
cent. ; several times the percentage was 30 or over, while, on the other hand, of
more than 5,000 eggs collected between July and October 1917 only 5 per cent.
produced Antestia. Of the 16,531 eggs collected during the period indicated, 12,882,
or 78 per cent., were parasitised, the average of the monthly percentages being
practically the same as this figure, namely, 77 per cent. ; the lowest monthly per-
centage was 54 per cent., the highest more than 90 per cent. The average monthly
percentage of eggs which did not go blue and did not hatch was 8 per cent., and
of those eggs which: did go blue 24 per cent. did not produce parasites. Of coffee
bug eggs laid in the laboratory and protected from parasites a small proportion
did not hatch. Some eggs, too, exposed to parasites in the laboratory turned blue
but did not produce parasites.

The causes of eggs not hatching have not been inquired into. Superparasitism
and hyperparasitism, which may quite likely be connected with the failure of
parasitised eggs to produce parasites, have not been investigated.

In the 9,750 parasites that were reared, the proportions of the sexes of the two

species were. as follows :—
S$ %SS LP WLP — Total.
Hadronotus ys hs 1168 22°6 4009 77-4 5177
Telenomus a cA 1079 23-6 3494 76°4 4573
For both species, therefore, the proportion of females to males was a little
greater than 3 tc 1. Figures from smaller collections from other plantations also
point to a ratio of 3 or 4 to 1.

The total numbers of the two species have just been given. These give as
percentages for the two species :—:-Hadronotus, 53 per cent. ; Telenomus, 47 per cent.
But it must be noted that at those periods when coffee bug eggs were most numerous
Hadronotus usually considerably outnumbered Telenomus. So that, if we take the
average of the quarterly percentages (see Table I) we then get :— Hadronotus, 43 per
cent. ; ZYelenomus, 57 per cent. Both these pairs of figures, however, indicate that,
over a long period, the numbers of the two species in this Kabete field were fairly
evenly balanced.

The variations in the frequency of the two species, for quarterly periods, are
shown in Table I (p. 200). We may note :—

1. The variations are considerable ; sometimes 70 per cent. or 80 per cent.
were Hadronotus, sometimes 80 per cent. or 90 per cent. were Telenomus.

194 F. W. DRY.

2. So far as the data go, the periods of ascendency of either species lasted for
from six to nine months. This time is sufficient, as the life-history figures will show,
for from four to six successive generations of parasites.

3. Ascendency of Hadyvonotus is associated with comparative abundance of
coffee bug eggs. Though an accurate index of the comparative abundance of eggs
was not obtained, it can be stated that they were most numerous from July to
October 1917, and the period of next greatest abundance was October 1918 to
March 1919. At the end ef 1917 and for the first three months of 1918 they were
very scarce indeed. During the remainder of the time they were moderately plentiful,
though not very numerous.

V. COLLECTIONS FROM OTHER PLANTATIONS.

The data as yet obtained are far too few for us to be able to correlate the occurrence
and subsidence of outbreaks of Antestia with parasite conditions, but the following
points may be mentioned :—

In collections of eggs made during outbreaks of Amtestia the percentages of
normal eggs have sometimes been high and the percentages of parasitised eggs
low. The following are examples :—

Percentage

not going

Total Percentage |Percentage| blue and
Place. Date. number | producing |parasitised. not

eggs. A ntestia. producing

anything.
1. | Near Nairobi ..| March 1918 ap 640 52 |) 46 2
2. | Limuru a ie Vay, LOS re ye 1008 58 37 5
3. | Limuru hs 2 «|| May 1920" *. a 1327 34 63 3

On the other hand, collections of eggs made during an outbreak of Antestia have
sometimes given a low percentage of normal eggs and a high one of parasitised eggs.
Such cases are :—

The Kyambu case numbered (4) is an example of an outbreak of Antestia subsiding
without any control measures having been taken by the planter. At the time the
collection was made the bugs were extremely numerous in part of the plantation.
In such cases it is usual to have the bugs collected by hand, but here, although they |
were left undisturbed, the following April the planter reported that the outbreak
had completely subsided. This was probably due to the work of the parasites.

The Kabete field in 1917 may also be cited. In June and July the bugs were
quite plentiful, In the months July to October 1917, as has already been stated

THE EGG PARASITES OF THE COFFEE BUG IN KENYA COLONY 195

only 5 per cent. of the eggs collected produced Antestia, more than 85 per cent.
being parasitised. Since then—up to the end of 1920—the bugs have never been
at all plentiful there.

The parasites are sometimes present in force in plantations where there has never
been a coffee bug outbreak. The following figures are for a field of coffee about
three years old, when the collection of eggs was made where there had never been
an outbreak :

Percentage
not going
Total Percentage | p, t blue and
Place. Date. number | producing aie a not produc-
eggs. Antestia, |PAtasiisec.) ing any-
thing.
7. | Near Nairobi Sept. 1918 re 862 4 86 10

The usual state of affairs in coffee plantations, it may here be remarked, is that
Antestia exists in small numbers only, but if the bug was present in a plantation,
parasitised eggs have always been found when a search has been made for them.

There are several cases on record when one outbreak of Antestia that has been
controlled by collecting the bugs by hand has been followed by another after some
such period as two years.

From some collections, notably from the Limuru district, Hadronotus has been
completely absent, though a very small number of Hadvonotus have been reared
from eggs from one Limuru plantation. Telenomus has been reared from every
collection examined.

VI. LIFE-HISTORY WORK ON THE PARASITES.

Many of the facts here given have already been published in Mr. Anderson’s
bulletin.

A. Hadronetus antestiae.
(1) Length of Life of Parasites.

For parasites kept in tubes closed with a pad of dry cotton-wool, provided with
water on a pad of cotton-wool moistened daily, and with coffee bug eggs in the
tube, the average length of life for either sex was about six days.

Five unmated females kept singly in tubes, provided daily with diluted golden
syrup on a pad of cotton-wool and supplied with coffee bug eggs, lived on the average
13 days. Further data for parasites which did not have access to coffee bug eggs
have given a slightly shorter time. The longest life of a female recorded is 16 days,
that of a male 13 days.

(2) Life-cycle of Parasites; Mated Females given Water only.
A series of experiments, in which the conditions defined below were adhered to,

was designed to compare the reproductive powers of the two species. The facts
for Telenomus will be given when the life-cycle work on that species is described.

The conditions were as follows :—(a) The experiment was set going not earlier
than 22.xi.17 and not later than 21.ii.18 ; (6) all parasites used as parents had emerged
from the coffee bug egg on the day on which the experiment was started ; (c) all the
parasites used were known to be virgin, as they had emerged in tubes not containing
any individuals of the opposite sex ; (d) the act of copulation was observed to take
place ; (e) once a parasite, male or female, had mated, no individual except its mate
was allowed access to it during the remainder of its life ; (f) the parasites were given

196 F. W. DRY.

no food, but a pad of cotton-wool was kept moist in the glass tube, which was closed
with a dry pad of cotton-wool; (g) the Antestza eggs used had been deposited in the
laboratory, and so kept as to prevent their accidental parasitisation ; () none of the
eggs showed eye-spots when given to the parasites ; (2) the number of eggs in each
experiment was 50; when the experiment started eggs to this number were placed
in a little cork tray having a rough floor so that the eggs would not roll about.

The results obtained from a series of ten experiments in which the above conditions

were observed were as follows :—

(a) The average lifetime of the parent parasites was 6 days, 8 days being the
maximum.

(6) On the average a little longer than 10 days elapsed before the first egg became
blue, and between 13 and 14 days before the last one to become blue had
changed ; it will thus be noticed that the time from parasitisation of the
egg to its going blue was about a third of the time from parasitisation to
emergence of the parasite.

(d) The total number of offspring obtained from the ten females was 233, or an
average of a little over 23 per female ; 256 eggs went blue, but 23 did not
produce parasites ; the highest number of offspring from one female was 34.

(e) Of the 233 parasites thus obtained the numbers of the sexes were :—Male, 43 ;
female, 190.

(3) Proportions of the Sexes.

At the same time as the series just described other experiments with mated
females, not fed but given water, were carried out in which one or more of the con-
ditions were not the same as those defined above. Particulars of some of these will
be given directly. The times obtained were not dissimilar from those just recorded,
and need not be given, but these other experiments give a larger number of parasites

reared in the laboratory from mated parents from which to calculate the proportions
of the sexes.

In 41 experiments, including the ten of the series just described, the numbers
of the sexes were :—Male, 178; female, 696. The percentages were thus almost
exactly :—Male, 20 per cent.; female, 80 per cent.

In every one of these 41 experiments the number of females exceeded the number
of males. In one experiment only all the offspring, 34 in number, were female.
In that experiment the parent male was the offspring of a virgin female. In six
other experiments in which the parent males were bred from virgin mothers the
offspring, totalling 31 males and 89 females, always included both sexes.

(4) One Male mating with several Females.

It was found that males will readily mate with several females. Mating, for
both species of parasites, is an affair of seconds. Occasionally a male would mate
twice with one female in rapid succession, but very often, and this applies also to
Telenomus, I noticed that a female which had been mated would run away from or
resist further attentions from the male.

One Hadronotus male, which emerged from a coffee bug egg on 30.xii.1917 and
was never given food or drink, and died on 4.1.1918, mated during the course of his
life with 16 virgin females. These 16 were all that were available ; with the first
two he mated twice, with the others once. These females were kept away from all
other males for the remainder of their lives, being kept singly in tubes, provided with
water, and supplied with eggs ; 369 parasites were in all bred from them, 68 male,

and 301 female. In every instance the number of females exceeded the number of
males,

THE EGG PARASITES OF THE COFFEE BUG IN KENYA COLONY. 197

This experiment, it may be remarked, gives us the evidence of 14 cases that a
single mating sufficed to produce a similar proportion of females to that found in
parasites reared from eggs from the field.

(5) Parthenogenetic Reproduction.

In a series of 20 experiments females to which no male had ever had access were
given coffee bug eggs. All the conditions defined for the series of ten mated pairs,
other than those relating to mating, were observed, and the times of the life-cycle
were not different from those obtained in that series. From these 20 females 442
offspring were obtained, all males; 482 eggs went blue, but 40 did not produce
parasites. The largest number from one female was 40. In other experiments
described under (6) and (7), in some of which the parent parasites were fed and the
number of offspring was larger, all the parasites reared were males. In no single
case under observation has a female been produced parthenogenetically.

(6) Parasitisation of Eye-spotted Eggs.

In two experiments with unmated females all the conditions of the series described
above under (2) were fulfilled except one ()—for all the 50 eggs, which had been
laid in the laboratory and protected from parasites, showed eye-spots when they
were given to the parasites. In these two experiments 28 parasites were bred, thus
showing that eye-spotted eggs can be parasitised. Just how late such eggs can be
successfully attacked by the parasites has not been determined.

When eggs are parasitised which do not show eye-spots, the change of colour
from white to blue is fairly sudden. Often on one day such an egg will be of the
typical white colour, and on the next of the typical blue. But when eye-spotted
eggs are parasitised the change of colour is more gradual. After a few days—a shorter
time than that in which eggs not eye-spotted would require to go blue—the colour
becomes a dirty brown, which deepens with successive days. In these two experi-
ments the eye-spots gradually faded and eventually could not be distinguished.
The eggs then took on the typical blue colour, but it was not so easy as with eggs
not eye-spotted to say exactly when the change took place.

(7) Number of Eggs parasitised by fed Females, unmated.

In each of five experiments one unmated female was given thirty different A ntestia
eggs every three days. The average number of offspring parasites, all males, obtained
in these experiments was 51, the maximum being 83. These numbers are considerably
higher than those obtained under the conditions of the experiments described above.

(8) Variations in the Length of the Life-cycle.

In hot weather the life-cycle is passed through more quickly than when it is
cooler. Particulars are given in Table II (p. 201) of a series of experiments in which
Antestia eggs, laid in the laboratory and protected from parasitisation, were placed for
a short time in a tube containing numerous parasites. The minimum times for the
life-cycle in the different experiments varied from 25 to 62 days. For meteorological
data, see Table IV (p. 201).

It will be noticed that the time elapsing before the eggs go blue is always about
a third of the period of the parasite’s life-cycle within the egg.

B. Telenomus truncativentris.

The facts obtained in life-history work on Telenomus will be discussed in much
the same order as for Hadvonotus. Unless stated otherwise, the conditions in
corresponding experiments on the two species were the same.

(1) Length of Life of Parasites.

The average length of life, for both males and females, was about 4 days when
they were given water only. When fed on diluted golden syrup the average length

198 F. W. DRY.

of life of unmated females provided with Antestia eggs was 22 days. Further data
for fed parasites of both sexes, some with access to Antestia eggs, some without,
have given a slightly less time. The greatest length of life of a female—one of those
provided with eggs—was 51 days, that of a male 16 days.

(2) Life-cycle of Parasites; Mated Females given Water only.

The results of a series of ten experiments corresponding to those described for
Hadronotus under A (2) were :—

(a) The average lifetime of the parent parasites, as already stated, was about
4 days, 6 days being the maximum.

(b) The first egg became blue after an average of 9 days, the last one after an
average of 11 days.

(c) As with Hadvonotus, the time from parasitisation of the egg to its going blue
was about a third of the length of the development within the coffee bug egg.

(d) The average time between the mating of the parasites and the first hatching
of their progeny was between 27 and 28 days.

(e) The total number of offspring obtained from ten females was 75, or 7) per
female ; 89 eggs went blue, but 14 did not produce parasites. The highest
number of offspring was 12.

(f) Of the 75 parasites thus obtained the numbers of the sexes were :—Male, 46 ;
female, 29.

(3) Proportion of the Sexes.

In the series of experiments of which the results have just been given considerably
more males were bred than females.

Like Hadronotus, Telenomus can reproduce parthenogenetically, all the offspring
so produced also being males. In the experiments under discussion, however, the
act of copulation was in every case watched. In eight of the experiments one or
more females were produced ; in two cases all the offspring were males, numbering
2 in one and 9 in the other. Now it has been shown that from collections of eggs
from the field, for Telenomus, as for Hadvonotus, the proportion of females to males
is about 3o0r4 tol. This, therefore, is the proportion of the sexes we should look for
in normal bisexual breeding. We therefore want to know—and this problem has
not been solved yet—why the proportion of females bred in the laboratory was lower
than that of females reared from eggs from the field.

A limited number of experiments with mated females fed on dilute golden syrup
were carried out, and the following short series may be taken as a hint which needs
to be followed up. It so happened that three females emerged from coffee bug eggs
in the presence of males about a day older than themselves. After all these parasites
had been together about a day they were separated into three pairs of male and female.
Each pair was put in a tube, given 30 Antestia eggs, and provided with dilute golden
syrup until death. The act of copulation was not observed. From these three
females the offspring bred were :—Male, 10; female, 40.

(4) One Male mating with several Females.

Males in the laboratory have mated with two, three, or four females. In no

experiments were males given the opportunity to mate with any number much larger
than four.

(5) Parthenogenetic Reproduction.

In a series of 20 experiments with unmated females the times of the life-cycle
were similar to those for mated females; 159 parasites, all males, were bred from
the 20 females ; 196 eggs went blue, but 37 did not produce parasites ; the largest
number from one female was 19. In other experiments to be described under (6)

THE EGG PARASITES OF THE COFFEE BUG IN KENYA COLONY. 199

and (7), in some of which the parent parasites were fed and the number of offspring
was larger, all the parasites reared were males. In no single case under observation
has a female been produced parthenogenetically.

(6) Parasitisation of Evye-spotted Eggs.

In one experiment similar to those described under A (6) 4 male Telenomus were
bred in eggs eye-spotted when they were offered to the parasite. The note about
the change of colour of eye-spotted eggs parasitised by Hadronotus is applicable
to those parasitised by Telenomus.

(7) Number of Eggs parasitised by fed Females, unmated.
In a series of five experiments with unmated females like those described for
Hadronotus the average number of offspring parasites, all males, was 22, the maximum

being 41. These numbers are, it will be seen, appreciably higher than those obtained
when the parasites were only given water.

(8) Variations in the Length of the Life-cycle.

As with Hadronotus, the life-cycle is passed through more quickly in hotter than
in cooler weather. Table III for Telenomus corresponds to Table II for the other
species. The minimum times for the life-cycle in the different experiments varied

from 46 to 21 days. Again, the eggs go blue after about a third of the period of
the parasite’s development within the egg has been passed through.

C. A Gomparison of the Data for Hadronotus and Telenomus.

As has been shown, many of the facts recorded for the two species are similar.
The chief differences to be noted are :—

1. In laboratory conditions, whether the parasites were fed on dilute golden
syrup or only given water, Hadronotus produced more offspring than Telenomus.

2. In the offspring of Hadronotus mated in the laboratory there was an excess
of females like that found in rearing parasites from the field. From mated
Lelenomus in laboratory conditions more males than females were generally

bred.

3. For a given time of year Telenomus passes through its life-cycle rather more
quickly than Hadronotus.

4. With hotter weather the life-cycle of Hadronotus is speeded up proportionately
more than that of Telenomus. (Contrast Tables II and III. Temperature
data are given in Table IV.)

VII. THE BEARING OF LABORATORY DaTA ON FIELD DATA.

Much more laboratory work is needed to elucidate the facts from the field, but
the following conclusions or suggestions may be put down.

1. On comparing the lengths of the life-cycles and the numbers of offspring per
female of Antestia and Hadronotus, and Antestia and Telenomus, we see that both
species of parasites have a quicker rate of reproduction than Antestia. But it must
be borne in mind that the egg-laying powers of Antestia are considerable. The
average number of eggs per female in the laboratory was 126. Nothing is known
of the mortality of nymphal coffee bugs in field conditions, so it is clearly possible
that when only some apparently small percentage—say 10 per cent.—of eggs are
producing Axdtestia, the numbers of the bug may be well on the increase.

2. In the field, with both species of parasites, mating is the rule ; parthenogenesis
is at any rate rare.

200 F. W. DRY.

3. Not so much light as could be desired is thrown on the problem of the vicissi-
tudes of the two species of parasites in the Kabete field. When we find two species
of parasites both attacking the same host we naturally ask why one does not succeed
in crowding out the other completely. As it is very unlikely that the resultant
of their reactions to the same set of conditions will be precisely the same, we are
disposed to look for some condition, not always constant, which affects the two
differently.

Such a condition we did find when it was learnt that the life-cycle of Hadronotus
is speeded up more in hotter weather than that of Telenomus. This, however, does
not give us the explanation of the facts from the Kabete field. There it appears
that comparative abundance of Antestia eggs results in the proportion of Hadronotus
going up, while scarcity of Antestia eggs would seem to send the proportion down.
We may make the obvious suggestion that under natural conditions, as in the
laboratory, Hadronotus has greater reproductive powers than Telenomus, but that
for some reason Telenomus is better able than Hadronotus to find the eggs in times
of scarcity. We need, therefore, to know much more about our two species.

With regard to the effects of temperature, reference may again be made to the
almost complete absence of Hadronotus from the several lots of parasites reared
from Limuru. The Limuru district, being higher than the Kabete field, is colder,
and it may be that it is this condition that gives the advantage to Telenomus.

VIII. CONCLUSION.

The facts so far ascertained have been recorded, together with some hints or
suggestions, and this account will have indicated a number of points calling for
further inquiry. Reference may be made to the life-history of the parasites within
the coffee bug egg, to superparasitism and hyperparasitism, to the effect of laboratory
conditions upon the proportions of the sexes of Telenomus, to the behaviour of the
parasites, and to a systematic following up of the parasites in a number of plantations.
These are some of the lines of work which need to be undertaken to enable us to
understand the interactions between Antestia, Hadronotus, and Telenomus.

TABLE I.

Collections of Coffee Bug Eggs from 6-acre Field, Government Farm, Kabete.

ena
\

Months during Total Hadronotus. | Telenomus.
PERIOD. which eggs were number of
collected. arasites.
iy e Noe of, No. %

1917.
3rd Quarter .. ae .. | July, Aug., Sept. ae as 2557 1469 57 1088 43
4th Quarter .. a6 toni} Oct. a 6 a A 955 627 66 328 34

1918.
Ist Quarter .. te ate 50 5 Sic ae De Eggs | very scarce.
2nd Quarter .. Re .. | April, June St ae oh 413 32 8 381 92
3rd Quarter .. bie .. | July, Aug., Sept. She a eh 219 28 552 72
4th Quarter .. ae se) | Oct: Nov. .Dec. fe Ne 1227 924 75 303 25

1919. a
Ist. Quarter’ ~. ae .. | Jan., Feb., Mar. ets ate 1579 1167 74 412 26
2nd Quarter .. Bye .. | April, May, June ao te 488 412 84 76 16
3rd Quarter .. aid ere te Me ats ure ac Eggs | not collected.
4th Quarter .. a .. | Nov. a3 ae as aud 340 153 45 187 55

1920.
ist. Quarter =< at ae. |eans ao a¢ ai nic 632 104 16 528 84
2nd Quarter .. ke .. | May nC ae ae cts 295 35 12 260 88
3rd Quarter .. re Je |) July; Sept. Ac ore sp 493 35 7 458 93
Totals e we te a6 9750 5177 53 4573 47 |
Averages of monthly percentages até af 43 oe 57

a EEE EES

THE EGG PARASITES OF THE COFFEE BUG IN KENYA COLONY. 201

TABLE II.
Variations in the Length of the Life-cycle of Hadronotus.

Antestia eggs exposed e
to parasites. va be) coir! Ones
wo we YO. epee] A 2 Hd
od 3 og a od og As
Hoe Efe! wood weg Oo BG
Ori 2 oO 095 ov of As
ora “4 Op PQ hp QWid oid
From. To. gee eaten aes gE5s 38

iS} ahi Se i= a)
Z32 SE 238 Za88 as
11.v11.18 12.vi1.18 21 11.1x.18 62 3 10
16.vi11.18 17.v11.18 20 14.1x.18 60 6 18
23.v11.18 24.v11.18 19 18.1x.18 57 4 20
31.v11.18 L.vur.18 21 24.1x.18 55 5 7] 8
4.vi1.18 5.vi11.18 20 24.1x.18 51 8 11
9.v111.18 10.vi11.18 19 26.1x.18 48 7 8
2.1x.18 3.1x.18 13 8.x.18 36 10 18
16.1x.18 We o.ent) 10 18.x.18 32 3 30
19.1x.18 20.1x.18 10 19.x.18 30 5 30
20.x11.18 23.x11.18 8 22.1.19 33 4 23
30.1.19 2.11.19 7 23.11.19 24 2 28
2.11.19 5.11.19 di 27.11.19 25 2 17

Particulars of the life-cycle during the months November 1917 to March 1918 are given

under VI.A.
TABLE III.
Variations in the Length of the Life-cycle of Telenomus.

rrr cr rN apres

wn
Antestia eggs exposed | a nu
cy |

to parasites. wien a % ; ears ar
Od | A gears) meee gs
Boe oy Hod HE ee = 4
ass $é | 82s woe :
Prom.) To. Ao» eee goku a2 5 se

| Bat Hoe eae: aoes °
Avo BAS ZU6 2368 BS
10.vi1.18 11.vir.18 13 25.v111.18 46 2 13
$2.vi11.18 23.v11.18 13 5.1x.18 45 4 36
23.v11.18 | 24.v11.18 13 5.1x.18 44 5 23
27.v11.18 | 28.v11.18 13 | 8.1x.18 43 1 9
4.vu1.18 | 5.v111.18 13 | 13.1x.18 40 3 21
9.v111.18 10.v111.18 14 | 18.1x.18 40 2 6
13.1x.18 14.1x.18 | 9 | 11.x.18 28 2 4
19.1x.18 20.1x.18 ) | 18.x.18 29 — 3
17.x11.18 20.x11.18 9 | 16.1.19 30 3 7
25.1.19 28.1.19 8 16.11.19 22 2 8
28.1.19 31.1.19 3) 18.11.19 21 cane 6
31.1.19 | 3.11.19 7 22.11.19 22 5 8

Particulars of the life-cycle during the months November 1917 to March 1918 are given
under V1.B.
SUN: 00 AV
Monthly Temperature Data, Government Farm, Kabete, 1917 to 1920.

1917. ; 191S* ; 1919. 1920.
Py” al ni) cad
fp o> | oe | 2
Month. Sid | Et 3 E<g Sic
2 2 eae = Ste |e = Seer | ae > |sus
3 = See 08 ft Ayes salad ies ees | aa @. |<od
ros Tea i * o q g * : 4 eS *K
oo ae Sea ok Ale SES > & of | DES ok og | DES
ne we GSA op 2 oe a= boo roa 83a oe op 3S
a8 a8 58” a o8& | 387 3 & of | a8" a o8 | 58
pops} ues Leste pilam wo ite A hfe fr uo bs pail ued mae bifesy
oe o% Bigt oe Otol aly Siete oe o* fas oe o4 Alain
canal > S OW Ss pa aS Oda Pane 2S OWS Po mS Om se
ts te OAD tz te OAD te te Oad Le aS OSs
| i |
Jen call = a) 72 61 46 78 62 52 a \ EE! 49 78 634
Feb. aye 32 Ae } 6224 48 Tks) 634 55 78 664 49 81 65
March .. 51 75 | 63 49 80 644 55 78 664 55 72 633
PeXgoymll 7) 55 71 63 54 73 634 56 74 65 56 73 644
May ..| 53 zi 62 53 70 614 54 72 63 58 61 594
ajime: ess 50 68 | 99 52 68 60 50 70 60 51 65 58
Jj se 47 69 58 50 68 39) 50 66 58 50 67 584
AUIS 2 50 70 60 | 49 70 594 48 72 60 49 69 59
Sept. <<. 49 73 61 49 75 62 52 74 63 47 73 60
Oct. bic 52 74 | 63 54 77 654 54 dee 63 52 73 624
INOVa! he 51 73 62 53 73 63 52 73 624 55 7A 63
Dees i 46 77 614 53 73 63 49 79; 62 53 70 614
— EEE — EE ee ee a ee ee ee ee

i Sr
a MAE PR
eee ee |
ae eae

y ie @ 7
i eh im

203

COREECTIONS RECEIVED.

The following collections were received by the Imperial Bureau of Entomology
between Ist April and 30th June 1921, and the thanks of the Managing Committee
are tendered to the contributors for their kind assistance :—

Mr. E. BatLarp, Government Entomologist :—60 Rhynchota ; from South India.

Mr. H. A. Battou, Entomologist, Imperial Department of Agriculture :—
11 Weevils and 8 early stages ; from Grenada.

Mr. C. F. C. BEEson, Forest Zoologist :—252 Coleoptera ; from India.

Mr. G. E. Bopxry, Government Economic Biologist :—13 Culicidae, 5 Hippo-
boscidae, 4 Siphonaptera, 2 Hymenoptera, 33 Coleoptera, 1 species of Coccidae,
10 other Rhynchota, 3 Orthoptera, 2 Mantispids, 5 Odonata, and 15 Mallophaga ;
from British Guiana.

Mr. Joun R. BoveEtt, Superintendent of Agriculture :—1 Culicid, 4 other Diptera,
8 Chalcididae, 5 Coleoptera, 6 Lepidoptera, 37 Isoptera, 2 species of Coccidae,
4 Aphididae, and 1 Pentatomid bug ; from Barbados.

Prof. C. K. Brain :—34 Coleoptera, 2 Trichoptera, 7 Planipennia, and 7 Odonata ;
from South Africa.

Major-General Sir Davip Bruce, K.C.B., F.R.S. :—100 Ants and 2 Weevils ;
from Madeira.

Mr. P. A. Buxton :—10 Culicidae, 27 Tabanidae, 5 Stomoxys, 95 other Diptera,
1 Dipterous pupa case, 1 Vespid, 107 Coleoptera, 4 Lepidoptera, 1 species of Coccidae,
19 other Rhynchota, 64 Orthoptera and 1 Dragon-fly ; from Palestine.

Director of Agriculture, Baghdad :—150 Hymenoptera, 7 Coleoptera, 50 Thysan-
optera, 4 species of Aleurodidae, 5 Orthoptera, and 100 Mites ; from Mesopotamia.

Director of Agriculture, N. Nigeria:—19 Hymenoptera and 7 Moths: from
West Africa.

Division of Entomology, Pretoria :—40 Coleoptera, 2 Lepidoptera, 1 Tingidid,
and 451 Orthoptera ; from South Africa.

Dr. H. Lynpuurst DuKE :—225 Isoptera, 1 Worm attacking Glossina, and 1 tube
of Fungi; from Uganda.

Mr. J. C. FaurE :—4 Rhynchota and 213 Orthoptera ; from South Africa.

Mr. T. BAINBRIGGE FLETCHER, Imperial Entomologist :—99 species of Coccidae ;
from India.

Mr. C. C. GowpEy, Government Entomologist :—2 Diptera, 23 Hymenoptera,
19 Coleoptera, 19 Lepidoptera, 30 Isoptera, 35 Thysanoptera, 8 species of Coccidae,
' number of Aphididae, 11 other Rhynchota, 2 Orthoptera, and 13 Ticks ; from

amaica.

Mr. C. B. HARDENBERG, Chief Entomologist, Department of Agriculture -—204
Coleoptera ; from Portuguese East Africa.

Mr. H. HARGREAVES, Government Entomologist :—34 Diptera, 41 Hymenoptera,
96 Coleoptera, 21 Lepidoptera, 2 species of Coccidae, 22 other Rhynchota, 6 Orthoptera,
14 Spiders, 3 Centipedes, 15 Crustacea, 9 Millipedes, and 20 Mollusca ; from Uganda.

204 COLLECTIONS RECEIVED.

Mr. G. F. Hitt, Entomologist, Australian Institute of Tropical Medicine :—
124 Culicidae, 52 Tabanidae, 5 Hymenoptera, 31 Coleoptera, 27 Lepidoptera, 9
Isoptera, 2 species of Coccidae, 13 other Rhynchota and 2 Orthoptera ; from Australia

Mr. 'M. Arzat Husain, Government Entomologist :—151 Parasitic Hymenoptera
and 19 Jassids; from the Punjab.

Mr. J. C. Hutson :—230 Thysanoptera ; from Ceylon.

Mr. R. W. Jack, Chief Entomologist, Department of Agriculture, Rhodesia :—
3 Coleoptera ; from South Africa.

Dr. W. A. LAMBorN :—4,368 Culicidae, 1 Haematopota, 27 other Diptera, 772
Hymenoptera, 2 Coleoptera, 3 Lepidoptera, 9 Lepidopterous puparia parasitised by
Chalcids ; from the Federated Malay States.

Dr. Li. Lroyp :—2 Dipterous larvae from human intestine, 2 Coleoptera,
3 Lepidoptera, 15 Thysanoptera, 2 tubes containing Tetranychus, and 20 Mites ;
from Cheshunt, Hertfordshire.

Major W. F. M. Loucunan, R.A.M.C. :—200 Ceratopogoninae ; from British
Honduras.

Mr. N. C. E. MitterR :—1 species of Coccidae ; from Tanganyika Territory.
Prof. S. A. MoKRzHETSKI :—7 Coleoptera (pests of roses) ; from Bulgaria.

Mr. J. C. Moutton :—1 Celyphid fly, 11 Coleoptera, and 1 Pentatomid bug; from
Singapore.

Mr. A. W. J. Pomeroy, Government Entomologist :—244 Diptera, 282 Hymen-
optera, 316 Coleoptera, 59 Lepidoptera, 118 Rhynchota, and 9 Orthoptera ; from
Nigeria.

Rhodesia Museum :—175 Diptera, 150 Coleoptera, 14 Thysanoptera, 9 Rhynchota,
2 Mantispids, and 1 Chrysopa ; from South Africa.

Mr. L. E. Ropinson :—4 Scolytidae and their borings ; from Colombia.

Senhor A. F. DE SEABRA :—1 Hippoboscid, 100 other Diptera, 14 Hymenoptera,
117 Coleoptera, 40 Rhynchota, 2 Orthoptera, and 1 Spider; from San Thome.

Mr. H. W. Seimonps :—30 Diptera, 6 Hymenoptera, 3 Coleoptera, 8 Rhynchota
and 2 Chrysopa; from Fiji.

Dr. R. vAN SOMEREN :—1 Tipulid, 3 Hymenoptera, and 2 species of Coccidae ;
from Kenya Colony.

Mr. R. Swarnson-Hatt, F.L.S. :-—5 Coleoptera, from the Island of Principe ;
93 Coleoptera, from San Thomé; 26 Coleoptera and 11 early stages, from Portu-
guese Congo.

fr. F. W. Uricu :—12 Siphonaptera, 2 Hymenoptera, 70 Ticks, and 2 tubes
containing Mites; from Trinidad.

Mr. RoBert VeIrcH :—124 Coleoptera and 5 Hymenoptera ; from Fiji.

NOVEMBER, 1921.

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205

THE INFLUENCE OF DROUGHT UPON MOSQUITO LIFE IN SURREY.
By Matcotm E. MACGREGOR,

Wellcome Bureau of Scientific Research
(Wellcome Field Laboratory, Waisley, Surrey).

The unusual and prolonged drought that we have experienced this year has had
a marked effect upon the mosquitos. So much so, that while last year I found no
difficulty in obtaining any of the eleven species which so far have been found to exist
in this locality, yet for many weeks now it has been impossible to obtain any
but a few Anopheles maculipennis (adults), Anopheles bifurcatus (larvae), Culrcella
morsitans (larvae, pupae, and adults), and Culex pipiens (larvae, pupae and adults).

As I have been anxious to get specimens of the other species, which were common
around here last year, in order to continue certain observations, I have made prolonged
searches for them in the usual breeding-places, only to find that with most of the water
collections dried up none of the species required was obtainable. .

Anopheles bifurcatus.

Normal conditions.—As larvae and pupae; plentiful in slowly moving streams
throughout the year. As adults; fairly numerous in pigsties from end of March
to June, but unobtainable later.

Conditions during drought.—As larvae and pupae; unobtainable, except in an
abnormal breeding-place (well). As adults ; unobtainable since late in June.

Last year I found that large numbers of Anopheles bifurcatus larvae were to be
found in a certain stream at Ockham throughout the summer and winter of 1920 and
until the streams dried up in June of this year. The only permanent water in the
vicinity was a small lake in which no Anopheline larvae were discovered last year.

I thought it probable that the Anopheles bifurcatus females after the stream had
dried up would be forced to oviposit on the water of this lake, but repeated search has
failed to discover larvae in the lake even under the present conditions of water
shortage.

Soon after, having made an extensive search for Anopheline larvae along the banks
on all sides of the lake, I came across a disused well in an open field near-by. Its
top was covered by a loosely fitting lid, and the water in the well was about five feet
deep, standing at about ten feet below the well-mouth. When examined, the water
was found to be crowded with the larvae of Culicella morsitans, associated with which
were numerous larvae of Anopheles bifurcatus in all stages of growth. Such an
unusual situation for Anopheles bifurcatus larvae, moreover in water that was turbid
and foul, is surely an outcome of the scarcity of water and the loss of normal breeding-
places. It is nevertheless difficult to account for the females’ choice of this situation
instead of the water of the lake, around whose weed-protected bays the eggs might
be laid in security against attack by fish. Under normal conditions, Anopheles
-bifurcatus chooses water that is clear and cool on which to lay her eggs.

The well, however, is the only situation where Anopheles bifurcatus larvae have
been found, and the proportion of adults of this species which can be emerging at
present must be infinitesimal compared with the normal.

(4183) P8/170 1000 9/21 Harrow G75 Q

206 MALCOLM E. MACGREGOR.

Anopheles maculipennis.

Normal conditions.—As larvae and pupae; plentiful in farm ponds and small
lakes from end of March until October. As adults; very numerous in pigsties
thronghout the spring and summer months. Hibernating females common during the
winter months.

Conditions during drought.—As larvae and pupae ; numbers relatively reduced
owing to drying up of many normal breeding-places ; but in certain permanent small
lakes, actual numbers of larvae and pupae increased, probably on account of the
scarcity of available breeding-places. As adults ; plentiful but not actually numerous
in pigsties.

It appears from my observations locally that Anopheles maculipenmis is the
species least affected by the drought. It chooses as a favourite breeding-place the
shores of large and small natural ponds and lakes which are, of course, the last
waters to dry up in conditions of drought ; and this species is consequently one of
the last to experience difficulty in obtaining breeding-places.

I should mention that pigsties in this locality have been found to be the favourite
haunts of adult Anopheles maculipennis and Anopheles bifurcatus, and that these
mosquitos are rarely found in stables. This state of affairs, which is so very different
from that in other parts of the country, is, I think, due entirely to the fact that all
the stables which I have visited in this county are ventilated at the junction of the
wall and roof. This method ensures a very thorough ventilation, and the average
temperature in the stables is much lower than the average temperature in stables
where ventilation is afforded by a few windows. Furthermore, because of the archi-
tecture, the amount of light in the average stable in Surrey is much greater than that
found in the average stable in Kent for instance, where Anophelines frequent the
stables in large numbers. This is a matter of such significance that attention should
be drawn to the value of the design in providing an “ all-the-way-round ”’ ventilation
between the walls and roof, as preventing the accumulation of mosquitos in stables.

The local pigsties, on the other hand, are low-built and the contained atmosphere
is close and warm at all times of the year, so that very favourable conditions are
maintained for the accumulation of mosquitos when they seek a blood meal, and
also conditions favourable to the hibernating females during the winter.

Anopheles plumbeus.

Normal conditions.—As larvae and pupae ; foundin almost every water-containing
tree-hole in a variety of different kinds of trees throughout the summer and winter
months. As adults; seldom observed, but occasionally in association with
Ochlerotatus spp. they have shyly attacked me while I have remained quietly seated
in the woods.

Conditions during drought—As larvae and pupae ; unobtainable now for many
weeks (last captured about middle of May), as all water in tree-holes has completely
evaporated. As adults; none observed since middle of May.

Anopheles plumbeus in this locality is quite common as larvae in water-containing
tree-holes and, as usual, is often associated with the larvae of Finlaya geniculata.

Very few adultshave been observed, and the one or two that I have caught sight of
have eluded capture. I have been anxious to obtain adult females for some time,
so as to get ova for the completion of certain observations. It occurred to me that
possibly another way of obtaining ova would be carefully to clear of detritus one of
the tree-holes in which Anopheles plumbeus had been breeding, and to keep it filled
with water artifically. Unfortunately this plan did not succeed, as such hosts of
insects regarded it as a providential water-supply when water was difficult to obtain
that the entrance to the hole was constantly choked with a seething mass of thirsty
insects.

THE INFLUENCE OF DROUGHT UPON MOSQUITO LIFE IN SURREY. 207

Culex pipiens.

Normal conditions.—As larvae and pupae; abundant principally in water-
troughs, rain-water barrels and other artificial water collections. As adults ; abun-
dant in houses, cellars, stables, etc.

Conditions during drought.—As larvae and pupae; relative numbers reduced,
but actual numbers in available breeding-places often increased. As adults ; easily
found in considerable numbers, but not so numerous as in normal years.

Culex pipiens being a species that largely makes use of artificial water collections
used by man, it follows that often these collections are maintained by him in spite of
dry weather, and thus Culex pipiens is not at so great a loss to find breeding-grounds
as are the less domestic mosquitos.

Culicella morsitans.

Natural conditions.—As larvae and pupae ; commonly found in certain wayside
ditches and in rain-water barrels. As adults ; quite common in out-houses, stables,
cellars and outside latrines, but never found in large numbers at a time.

Conditions during drought.—As larvae and pupae ; scarce and now only found in
one rain barrel kept supplied with water artificially, and in one instance found in a
shallow well ten feet below ground. As adults ; none observed for many weeks.

Culicella fumipennis.

Normal conditions.—As larvae and pupae ; common in certain wayside ditches.
As adults ; frequently found on the walls in out-houses and stables.

Conditions during drought.—As larvae and pupae; unobtainable. As adults ;
unobtainable.

Finlaya geniculata.

Normal conditions.—As larvae and pupae; abundant in the water-containing
holes in a variety of trees. Often associated with the larvae and pupae of Anopheles
plumbeus. As adults ; sometimes observed to attack human beings while they are
in the woods, though not very often seen.

Conditions during drought.—As larvae and pupae ; unobtainable since middle of
June. As adults; unobtainable, and not observed in the woods since beginning
of June.

Soon after the tree-holes had begun to dry up generally, numerous larvae and
pupae of Finlava geniculata were collected from several holes that still contained
water, and these were brought to the laboratory for experiments on their develop-
ment in waters giving different reactions. Many were placed under conditions
that highly favoured development and emerged as large and active mosquitos.
They were at first kept in a cage in the laboratory and fed upon dates and human
blood in an attempt to obtain ova from them, but while all the females partook
of anample diet, fertilisation apparently would not take place under caged conditions,
even when there were about an equal number of active and healthy males in associa-
tion. In the hope of getting ova from them under natural conditions, the whole
batch of mosquitos was carefully captured in tubes and taken to the vicinity of
a beech-tree in which the tree-holes were kept full of water, and there released.
Possibly on account of the fact that the water in the holes was besieged by insects
of all kinds (see under Anopheles plumbeus), no eggs were laid, nor were any of the
released mosquitos seen again.

(4183) Q2

208 MALCOLM E. MACGREGOR.

It was observed in the laboratory that the “bite” of I’imlaya geniculata is
distinctly painful. The proboscis is large and the initial prick of the “bite”’ is
sharp, but unlike the usual conditions when other mosquitos bite (a painless period
following the initial prick when once the insect has got the proboscis well embedded
in the tissues and has begun to feed), the initial prick from Finlaya geniculata is
followed by a burning sensation, which continues and increases while the insect
feeds ; so much so that a good deal of will-power has to be exercised against the
impulse to jerk the mosquito off. The subsequent effects of the “ bite ’’ are not,
however, more serious than the usual effects of a mosquito bite.

Ochlerotatus caspius.

Normal conditions.—As larvae and pupae; found in numbers in a woodland
pool near Ockham. As adults ; not observed under natural conditions, but several
bred out from larvae in the laboratory.

Conditions during drought.—As larvae and pupae; unobtainable; pool com-
pletely dried up since last May. As adults; unobtainable.

Ochlerotatus nemorosus.

Normal conditions—As larvae and pupae; abundant in woodland pools and
streams. As adults ; very large numbers obtainable in the woods, where, particularly
in the afternoons, they attack so persistently that there is no peace to be had.

Conditions during drought.—As larvae and pupae ; unobtainable since the wood-
land pools dried up in May. As adults; unobtainable since about the middle of
June, when it was found that even in the afternoons one might frequent the woods
in peace, unmolested by a single mosquito—a state of affairs very different from
what obtained during the whole of the summer months last year.

Ochlerotatus waterhousei.

Normal conditions.—As larvae and pupae; numerous in certain woodland
pools. As adults; numerous in the woods where, associated with Ochlerotatus
nemorosus, they persistently attack human beings, particularly in the afternoons.

Conditions during drought.—As larvae and pupae; unobtainable since end of
May. As adults ; unobtainable since May.

Theobaldia annulata.

Normal conditions.—As larvae and pupae ; numerous in certain wayside ditches
and in rain-water barrels. As adults; common in stables and outhouses, though
never numerous.

Conditions during drought—As larvae and pupae; unobtainable. As adults ;
a few still obtainable in pigsties in association with Anopheles maculipenmis.

* * BS * * * *

From the foregoing records it will be seen that drought tends to affect the
mosquito population of an area adversely, decreasing the normal breeding-places,
and in some instances, in the case of particular species, causing the normal breeding-
places to disappear entirely. This is true for such species as Anopheles plumbeus,
Finlaya geniculata, Ochlerotatus caspius, Ochlerotatus nemorosus and Ochlerotatus
waterhouset. It will be interesting, therefore, to see whether these species will be

THE INFLUENCE OF DROUGHT UPON MOSQUITO LIFE IN SURREY. 209

rare in this locality during future years. Predictions were made by several people
at the beginning of the hot weather that there would probably be an enormous
increase in the number of mosquitos all over the country, and that in consequence,
with the large numbers of ex-soldiers in our midst who had suffered from malaria,
we should probably see many new cases of indigenous malaria. The reverse has
been true. I learn from the Ministry of Health that the number of cases of
indigenous malaria contracted in England in 1920 was 36; whereas in 1921, up to
the 29th August, only four cases had been recorded. Hot weather is obviously
only one of the many factors in the malaria equation. Even in England, where
it had been maintained that the historical epidemics of malaria have been occasioned
by a hot summer following the return of malaria-infected troops from abroad, it will
be clear that this combination is not all that is necessary to produce even a mild
epidemic of indigenous malaria.

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211
SOME COCCIDAE FROM EASTERN ASIA.

By G. F. Ferris,
Stanford University, California.

Through the kindness of Professor E. O. Essig, of the University of California,
I have been enabled to examine a collection of CoccrpAE from Formosa, made
by M. Maki and S. Inamura, and also a small amount of material from Foochow,
China, collected by Mr. C. R. Kellogg. It is upon this material that the present
paper is for the most part based. In addition to this, I am taking advantage of the
opportunity to rectify certain errors that have been made in the identification of
a few species from Japan.

The material from Formosa is especially interesting, as nothing appears to have
been published concerning the Coccid fauna of that island. The majority of the
species here recorded are widely-distributed forms ; a few are known from Japan ;
and several are species that have heretofore been recorded only from India, Ceylon
and Java. While the number of species dealt with is not sufficiently large to permit
of any generalisation, there is at least a suggestion that the Coccid fauna of Formosa
is more closely related to that of southern Asia than to that of Japan.

The types of the new species here described are in the Stanford University
Collection.

Icerya seychellarum (Westw.).
On Morus alba and Citrus sp., Taihoku, Formosa.

icerya purchasi (Maskell).
On Citrus sp., Taihoku, Formosa.

Pseudococcus filamentosus (CkIl.).
On Citrus sp., Taihoku, Formosa.

Pseudococcus comstocki (Kuwana).

On Citrus sp., Taihoku, Formosa. Previously recorded from Japan and the
United States.

Pseudococcus virgatus (Ckll.).
On Baphima sp., Ako, Formosa.

Pseudococcus citri (Risso).
On Morus alba, Ako, Formosa.

Antonina bambusae (Maskell).

On Bambusa stenostachya, Taihoku, Formosa. This is the species that has
ordinarily passed under the name of Chaetococcus bambusae. Iam entirely in accord
with Green in the opinion that Chaetococcus cannot be maintained as distinct from
Antonina.

Antonina crawii (Ckll.).
1902. Eriococcus graminis (2) Maskell ; Kuwana, Proc. Calif. Acad. Sci. (3), 3: 50.
On bamboo, Taihoku, Formosa. Kuwana (ref. cited) has recorded Evzococcus
gramunis, Maskell, from Japan. I have at hand the specimens upon which the record

was based and they prove to be nothing more than immature stages of an Antonina,
without much doubt 4A. crawit.

2, G. F. FERRIS.

Cerococcus ficoides (Green).

On Mallotus japonica, Taihoku, Formosa. Previously known only from the
original record, on tea from Bengal. The excellent description and figures given by
Green render the identification practically certain.

Tachardia decorella (Maskell).
On Ficus retusa and undetermined host, Taihoku, Formosa.

Mallococcus sinensis (Maskell).

On undetermined host, Foochow, China. I have at hand part of the type
material of this species and would redescribe it here (it is hardly or not at all
recognisable from the original description) were it not that another author has a
redescription now in press.

Ceroplastes rubens (Sign.).
On Citrus, Ako, Formosa.

Coccus bicruciatus (Green).
On Murraya exotica, Taihoku, Formosa. Previously recorded only from Ceylon.

Coccus elongatus (Sign.).
On Acacia confusa, Codiaeum variegatum, Gossypium herbaceum, Hibiscus
vosa-sinensis and Myrica rubra, at Taihoku, and on Morus alba at Ako, Formosa.

Saissetia hemisphaerica (Targ.).
On Gardenia florida, Taihoku, Formosa.

Fig. 1. Avlacaspis cinnamomi (Newstead): A, adultfemale; B, portion of pygidial margin ;
C, pygidium,
Aulacaspis cinnamomi (Newstead) (fig. 1).

On Cinnamomum camphora, Taihoku, Formosa. Previously recorded only from
Cinnamomum ceylanicum in Java.

SOME COCCIDAE FROM EASTERN ASIA. QS

The specimens at hand differ somewhat from the figures given by Newstead,
but these figures are not entirely clear, and in view of the close similarity in general
appearance and‘in hosts I am inclined to assume that the figures are in error. I am
presenting new figures. The species is certainly an Avlacaspis.

Aulacaspis rosae (Bouché).
On cultivated rose, Taihoku, Formosa.

Aulacaspis pentagona (Targ.).
On Morus alba, Taihoku, Formosa.

Fig. 2. Aulacaspis tegalensis (Zehntner): A, adult female; Bb, portion of pygidial margin ;
C, pygidium.

Aulacaspis tegalensis (Zehntner) (fig. 2).

On sugar-cane, Taihoku, Formosa. Previously recorded from Java and Mauritius.
My determination is on the basis of specimens from Java.

I know of no figures of this species other than those given by Zehntner, which
are in out-of-the-way publications and are in addition not especially satisfactory,
and as the species is likely to be of some economic importance I am_ presenting
new figures. JI regard it as an Aulacaspis rather than a Chionaspis.

Phenacaspis eugeniae (Maskell).
On Michelia fuscata, Taihoku, Formosa.

The exact identity of Maskell’s Chionaspis eugemiae appears to be in doubt, for
there occurs in the Pacific region a series of forms which are very closely related.
The Formosa specimens agree closely with examples from Hawaii which have been
determined as eugentae.

Chionaspis annandalei (Green).

On Bambusa stenostachya, Taihoku, Formosa. Previously known only from
India.

214 G. F. FERRIS.

Pinnaspis simplex, sp. n. (fig. 3).

From undetermined host, Foochow, China.

Scale of the female 1-5 mm. long, elongate, narrow, dark brown. Scale of the
male not observed.

Female 0-9 mm. long, of the usual elongate form, the derm membranous throughout
except for the pygidium. Margins of the abdominal segments projecting but little,
without gland spines or at the most with a single gland spine on the last segment
anterior to the pygidium, with numerous marginal ducts. Dorsum without ducts.
Anterior spiracles with a small group of pores. Pygidium (fig. 3) acutely pointed,
the median lobes alone developed, apparently fused, their outer edges crenulate.
Extending into the pygidium from the median lobes is a slender, elongate median
thickening. Beyond the median lobes there is first a small prominence followed
by two setae, one of which has a conspicuously chitinised socket, a gland spine, two
submarginal pores, a seta, a submarginal pore, two gland spines and a submarginal
pore. Dorsum of the pygidium without pores except for the marginal series and a
single pore representing the front row. Anal opening close to the anterior margin
of the pygidium. Circumgenital pores in five groups.

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Fig. 3. Pinnaspis simplex, sp.n.: pygidium and portion of pygidial margin.

Notes.—Specimens of this species have been examined by Mr. E. E. Green, who
agrees with me that it is undescribed. It is characterised by the single pair of lobes
and is quite distinct from any other species known to me. I may note that I am
considering the genera Pinnaspis and Hemichionaspis as synonymous.

Pinnaspis minor (Maskell).
On undetermined host, Taihoku, Formosa.

Parlatoria pergandei (CkIl.).
On palm (?), Foochow, China, and on Thea sinensis and undetermined host at
Taihoku, Formosa.

Parlatoria zizyphi (Lucas).
On Citrus sp., Taihoku, Formosa.

SOME COCCIDAE FROM EASTERN ASIA.

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Fiorinia japonica (Kuwana) (fig. 4, C).

1902. Fiorinia fioriniae var. japonica, Kuwana, Proc. Calif. Acad. Sci. (3),
<3: 79 (part).

From Pinus thunbergii, Taihoku, Formosa. Originally recorded from Pinus sp.
and Podocarpus chinensis in Japan, the latter record in error.

In the original description of this species as a variety of I’. fioriniae the type
host was not designated. I find the material recorded by Kuwana to include two
species and designate as a lectotype a specimen from Pinus. The specimens from
Podcarpus I consider to belong to F. juntpert, Leonardi.

Fig. 4. Pygidial margins of : A, Fuorvinia fioriniae (Targ.), from palm in the United

States; B, F. junipert, Leonardi, from Podocarpus chinensis in Japan; C, F. japonica,

Kuwana, from Pinus in Japan; E, F. chinensis, sp.n.; F, F. chinensis, sp. n., second stage.
D, pygidium of F. chinensis, sp. n.

Kuwana separated this form from F. forimiae on the grounds of the larger
number of circumgenital pores, but there appear to be more satisfactory characters in
the number and arrangement of the marginal tubular ducts of the pygidium. In
F. fioriniae (fig. 4, A) there are not more than four large ducts followed by as many
small ducts. Otherwise the two species are very similar.

Fiorinia juniperi (Leonardi) (fig. 4, B).

1902. Fiorinia fioriniae var. japonica, Kuwana, Proc. Calif. Acad. Sci. (3),
3:79 (part).

Specimens from Podocarpus chinensis in Japan were included by Kuwana under
F, fioriniae var. japonica, but I consider them to belong rather to F. junipert. They
agree very Closely with specimens of jumipert (det. Green) from Ceylon. The species,
like japonica, is very similar to fioriniae, but differs in having as many as eight large
marginal ducts instead of four on the pygidium (fig. 4, B).

216 G. F. FERRIS.

Fiorinia chinensis, sp. n. (fig. 4, D, E, F).
From undetermined host from China, taken in quarantine at San Francisco.
Scale not available for description.

Female 0:75 mm. long. Without a process between the bases of the antennae,
with no pores about the spiracles and with at the most a very few small gland spines
on the margins of the abdominal segments. Pygidium (fig. 4, D), with the median
lobes quite large, rounded, minutely serrate, connected at the base and bounding a
median notch in which are a pair of small gland spines. Second pair of lobes (fig. 4, E)
represented merely by a pair of small tooth-like projections. Between the first and
second lobes is the opening of a duct and beyond the second lobe are two more sub-
marginal ducts, all quite small. Circumgenital pores present, forming an almost
continuous arch.

Second stage without marginal projections on the abdomen, the pygidium (fig.
4, F) with a deep median notch, the median lobes narrower than in the adult, the
second pair represented by a single undivided lobe. Lateral margins of the pygidium
with several small gland spines and with five submarginal ducts.

Notes.—This species would appear to belong to Leonardi’s genus Trullifiorima,
but I am not at present inclined to accept his groups. It somewhat resembles
F. rubrolineata, Leon., but differs in the larger lobes and submarginal ducts.

Lepidosaphes beckii (Newman).
On Murraya exotica, Taihoku, Formosa.

Fig. 5. Lepidosaphes tubulorum, sp. n.: A, lateral margin of abdominal
segment; B, pygidium.
Lepidosaphes tubulorum, sp. n. (fig. 5).
1902. Mytilaspis pomorum (L.); Kuwana, Proc. Calif. Acad. Sci. (3), 3:80
(part).
Type from Sapium sebiferum, Taihoku, Formosa. Also from Salix warburgi at
the same place, and from Jlex crenata, willow and currant in Japan.

Scales of male and female of the type common to the genus, dark brown, the former
moderately broad, length 2-75-3 mm.

SOME COCCIDAE FROM EASTERN ASIA. pA)

Female 1-1 mm. long, elongate, rather broad ; the derm membranous except for
the pygidium ; the margins of the last three abdominal segments projecting and
bearing numerous gland spines and likewise bearing a small spur-like, chitinised pro-
cess (fig. 5, A). In some specimens this process may be developed on but part of
the segments. Dorsum of the abdomen with large numbers of extremely small
ducts.

Pygidium (fig. 5, B) with two pairs of lobes ; the median pair widely separated,
broad and with a deep subapical notch on each side, the second pair bilobed.
Between the median lobes is a pair of small gland spines ; between the median and
second lobes a pore prominence with two small projections ; beyond the second lobes
a gland spine, a pore prominence with two large pores, two gland spines, two large
marginal pores, two gland spines and two pores. Margins of the marginal pores
heavily chitinised. Dorsal ducts extremely minute, arranged in three irregular areas
somewhat variable in number. Circumgenital pores in five groups of 6-13 pores.

Notes.—The specimens recorded from Japan were included by Kuwana in Lepido-
saphes ulmi (=Mytilaspis pomorum) along with specimens really referable to that
species and others (from orchid} belonging to a species that I shall not here consider
because of the scantiness of the material. While L. tuwbulorum is indeed quite similar
to L. ulmi, it is readily separable by the small size and the arrangement of the dorsal
ducts.

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Fig. 6. Lepidosaphes japonica, Kuwana : pygidium.

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Lepidosaphes japonica (Kuwana) (fig. 6).

1902. Mytilaspis pomorum var. japonica, Kuwana, Proc. Calif. Acad. Sci. (3),
3: 80.

From Abies firma, Japan.

I have at hand the types and type material of this species. It cannot be regarded
as having any connection with L. ulmi, being in fact a very distinct species.
I present the following notes.

218 G. F. FERRIS.

Female with the derm unchitinised except for the pygidium. Margins of the
abdominal segments projecting, the last three with three or more gland spines, the
margins of both thorax and abdomen and the dorsum of the abdomen with numerous
moderately large ducts. Pygidium (fig. 6) with two pairs of lobes, the median pair
rounded, the second pair bilobed. Between the median lobes are two small gland
spines ; between the median and second lobes a pore prominence with two projecting
points; beyond the second lobes two small gland spines, two large submarginal
pores, two gland spines, two pores, three gland spines and a pore. Margins of the
marginal pores heavily chitinised. Dorsal ducts only slightly smaller than the
marginal ducts, arranged in three rows as indicated in the figure.

Genus Pygalataspis, nov.

Diaspidine CocciDAE referable by the character of the ducts to the Diasis series,
that is with the ducts short, relatively broad and with a pair of transverse bars across
the inner extremity. Ducts abundant on the pygidium, both dorsally and ventrally,
not arranged in rows. Gland spines of the ordinary simple type lacking, being
replaced by a series of short, broad, variously toothed plates along the margin of the
pygidium, these plates apparently arising in part from the margins of the lobes.
Two pairs of lobes or lobe-like processes present, the outer pair not bilobed. — Cir-
cumgenital pores present in five groups. Scale of both sexes elongate, with the
exuviae at one end, that of the female with the dorsal and ventral portions continuous,
the ventral scale composed in part of the ventral portion of the second exuviae.

Type of the genus, Pygalataspis miscant, sp. n.

Notes.—In the abundance and distribution of the ducts and the correlated charac-
ters of the scale this genus most closely resembles Odonaspis, but the peculiarly
shaped plates and the extraordinarily large lobes are quite unlike anything else with
which I am familiar. The genus Odonaspis is usually attached to the Aspidiotus
series of the DIAsPIDINAE, but I am inclined to doubt the correctness of this placing,
regarding it rather as belonging to the Diaspis series.

Pygalataspis miscanthi, sp. n. (fig. 7).
From Miscanthus sinensis (a grass), Taihoku, Formosa.

Scale of the female about 2-5 mm. long, elongate, white or brownish ; scale of the
male similar in form and colour to that of the female, about 1 mm. long.

Female (fig. 7, B) elongate, with nearly parallel sides, the margins of the abdominal
segments projecting but little or not at all, the derm membranous except for the pygi-
dium and the lateral margins of the last two or three abdominal segments which are
heavily chitinised. Lateral margins of the metathorax, and of all the abdominal
segments and the dorsum of the last two segments with numerous ducts of the type
shown in fig. 7, D.

Pygidium (fig. 7, A) with the marginal area heavily chitinised and more or Jess
folded. Two pairs of large, rounded lobes or lobe-like processes present, each with
a broad, flattened, irregularly toothed plate arising from the outer margin, the plates
of the second pair larger than those of the median pair. Beyond the second is a
cluster of two or three smaller plates. Setae at the bases of the lobes, both dorsally
and ventrally, long and slender. Tubular ducts very abundant, all small, those of
the venter confined to a broad marginal zone. Circumgenital pores in five groups of
15 or more pores.

Second stage with the pygidium (fig. 7, C) terminating in a pair of prominent,
pointed processes, which bear small plates as in the adult, and with a few small plates

SOME COCCIDAE FROM EASTERN ASIA. 219

along the lateral margin beyond the lobes. Ducts few, relatively large, confined to
a submarginal series. Second exuviation occurring by the splitting of the derm
about the posterior lateral margin, the ventral portion becoming incorporated in the
ventral scale.

Fig. 7. Pygalataspis miscanthi, sp. n.: A, pygidium; B, adult female; C, pygidial
margin of second stage; D, type of duct.

Odonaspis penicillata (Green).

1902. Aspidiotus inusitatus, Green ; Kuwana, Proc. Calif. Acad. Sci. (3), 3:65
(misidentification).

On Bambusa stenostachya, Taihoku, Formosa. Specimens recorded from Japan
as O. inusitata by Kuwana (ref. cited) are this species instead.

Aspidiotus lataniae (Sign.).
On Morus alba, Ako, Formosa, and undetermined host from Formosa in quaran-
tine at San Francisco.

Chrysomphalus aurantii (Maskell).
On Citrus sp., Taihoku, Formosa.

220 G. F. FERRIS.

Chrysomphalus aonidum (L.).
On Cycas revoluta, Ako, Formosa.

Pseudaonidia duplex (Ckll.).
On Michelia fuscata, Taihoku, Formosa.

Pseudaonidia trilobitiformis (Green).
On Citrus sp., Taihoku, Formosa.

THE, LARVAL AND PUPAL SPAGES ‘OFTHE’ BIBIONIDAE:

By Husert M. Morris, M.Sc., F.E.S.

Entomological Department, Institute of Plant Pathology, Rothamsted Experimental
Station, Harpenden.

Several species of BIBIONIDAE are very common in the adult state during the
spring and early summer in Britain, but their larvae seem to have been very little
studied. The latter frequently occur in large numbers together, and are from time
to time reported to have caused damage to various crops.

Probably the commonest species of Bibio are B. marci and B. johannis, but
several others are also very frequently met with. The life-history of Bibio johannis,
L., has been fully dealt with previously (6), and in the present paper it is proposed
to give some account of the life-histories of Bzbio marci, L., B. lacteipennis, Ztt.,
and B. venosus, Mg.

. I am greatly indebted to Dr. A. D. Imms for suggestions and advice during the
course of this work, and for the material of Bzbio venosus, and to Mr. F. W. Edwards
for identifying the adults which have been reared.

Bibio marci, L.
Oviposition.
In the early part of May 1920, a number of adults of Bibio marci, of. both
sexes, were captured and placed in a glass jar, the top of which was covered with gauze.

Fig. 1. Eggs of Bibio marci, x 6.

At the bottom of the jar a layer about 3 cm. in depth of fine damp soil was placed
The flies were fed on sweetened water, and by this means it was found possible to
induce the females to oviposit. After a day or two, the time probably varying
with their age when captured, females were seen making their way into the soil.
This is accomplished almost entirely by means of the fore legs, which are moved
up and down in front of the head, with their tarsi bent back, the actual pressing
back of the soil being done by the distal ends of the tibiae. In this manner a
burrow just large enough to contain the fly is made, the insect turning over from
time to time so that the soil is pressed aside in all directions. The burrows have a
very irregular course, but the flies gradually make their way downwards into
the soil.

At a little distance below the surface, the depth varying in different cases, a small
cell is constructed which is of greater diameter than the burrow, and in this cell a
mass of eggs is deposited, the eggs being arranged in an orderly manner (fig. 1).

(4183) R

222 HUBERT M. MORRIS.

The depth of soil in the jar was not sufficient to show the maximum depth to which

the flies would penetrate for the purpose of oviposition, as some of the egg-masses _

were in contact with the bottom of the jar, 3-5 cm. below the surface.

After oviposition the flies soon died, usually just outside the cell containing
the eggs.

Eggs.

The jar containing the soil in which the eggs had been laid was placed in the |
laboratory, the soil being kept damp. The eggs were observed to hatch in from |
33 to 35 days after being laid, all those in a mass hatching more or less at the —

same time.
The eggs are white when first laid, but soon afterwards darken at both ends.

They are almost cylindrical, with rounded extremities, the diameter of one end being |
slightly greater than that of the other, and they are slightly curved. Their length —
is from 0-73 mm. to 0-76 mm.; their diameter at the larger end 0-18 mm., and |

at the narrower end 0-15 mm.

The eggs appear rather opalescent owing to their being covered with small
projections on the chorion.

Fig. 2. Newly hatched larva of Bibio marci, lateral view ; setae of
right side omitted; x 65,

Newly hatched Larva.

The newly hatched larva (fig. 2) is about 1-6 mm. in total length, the length of
the head being about 0-25 mm. The head is relatively large, of a pale yellowish-
brown colour, and bears a number of relatively long setae.

The body is colourless and transparent, the contents of the alimentary canal
showing through as a dark line. The body is divided into twelve segments, each
bearing a number of long, slender setae. There is, in addition, a much greater
number of shorter setae, all the setae arising from enlarged conical bases.

At this stage the larva bears a single pair of spiracles, which are situated on the
twelfth segment, in a corresponding position to that occupied by the spiracles of that
segment in the full-grown larva.

The head and mouth-parts are very similar to those of the full-grown larva.
The mandibles are shorter, with relatively longer teeth; the maxillae are similar
to those of the older larva; the projections at the anterior end of the submentum
of the labium are sharper and relatively longer, and there is a median projection
between them, which is not present in that of the full-grown larva.

THE LARVAL AND PUPAL STAGES OF THE BIBIONIDAE. 223

Fully grown Larva.

Well grown larvae of Bibio marci were found in considerable numbers during
the winter amongst the decaying remains of grass which had been cut the year
before, or earlier, and allowed to rot on the ground. They were also found amongst
the roots of grass.

Fig. 3. Fully grown larva of Bibio marci, x 5; sp. 1, spiracle of first segment ;
sp. 12, spiracle of twelfth segment.

When fully grown the larva is from 20 to 24 mm. in length, and about 2-5 mm.
in breadth. Its thickness is rather less than its breadth. The length of the head
is about 1-5 mm. and its breadth a little less (fig. 3).

Fig. 4. Head of fully grown larva of Bibio marci, dorsal view, x 53;
a, antenna ; md, mandible ; mp, maxillary palp.

The head (figs. 4, 5) is dark brown and shining, appearing almost black ; the
anterior half is somewhat darker in colour than the posterior and bears several
long setae. The form of the mouth-parts and antennae is shown in figs. 6-9. The

(4183) a

224 HUBERT M. MORRIS.

antennae are exceedingly small and inconspicuous, being situated just above the
bases of the mandibles. The structures which were previously considered to be the
antennae (6) are lateral protuberances of the labrum.

Fig. 5. Head of fully grown larva of Bibio marct, ventral view» x 53; J, labium ;
md, mandible; mp, maxillary palp; mx, maxilla.

The body is nearly cylindrical, but somewhat flattened dorso-ventrally, and
normally is slightly curved, the ventral surface being concave. The body is divided
into twelve segments, of which the first is the longest. It bears ten pairs of spiracles,
which are situated a pair on each segment except the second and eleventh, and
project slightly from the body. The spiracles on the first segment are about twice
the size of those on segments 3 to 10, and those on segment 12 are about four times
the size of those on segments 3 to 10, and each one has two openings.

Fig. 6. Antenna of larva of
Bibio marci, X 580.

Each segment bears a number of stout conical processes, which are arranged
more or less in rows, on both the dorsal and ventral surfaces, the more lateral processes
in the rows being usually at least twice as long as the median ones. The arrange-
ment of these processes does not appear to vary in the different species of Bzbio
which have so far been examined.

THE LARVAL AND PUPAL STAGES OF THE BIBIONIDAE. 225

The spiracles of the first segment are situated laterally near the posterior boundary
of the segment. Dorsally the segment bears two rows of processes, each row
containing eight processes, the lateral processes of the second row being slightly

Fig. 7. Right mandible of larva of Fig. 8.
Bibio marci, ventral view, x 100.

Right maxilla of
larva of Bibio marci,
ventral view, x 100.

displaced forwards by the spiracle. Ventrally the segment bears three incomplete
tows. The first row consists of two processes ; the second of four, one pair near
the median line and the other pair lateral ; in the third row there are four processes.

The second segment, which has no spiracles, bears a row of eight processes dorsally
and another row of eight processes ventrally ; the median pair in the latter row
is situated nearer to the anterior margin of the segment than the remainder.

Fig. 9. Labium of larva
of Bibio marci, ventral
view, x 100.

The third segment bears a pair of spiracles laterally towards its anterior margin ;

dorsally it bears a row of eight processes, and ventrally a similar number, of which
the middle pair is placed forwards as on the second segment.

The fourth segment bears a pair of spiracles in a similar position to those on the
third segment. Dorsally this segment bears a row of eight processes, the lateral

226 HUBERT M. MORRIS.

pair being slightly nearer the anterior end than the remainder. On its ventral
surface this segment bears two rows of processes, the first row containing
six and the second four. The segments five to ten are similar to the fourth
segment.

On the dorsal surface of the eleventh segment there is a row of six large pro-
cesses near its posterior margin, of which the median pair is the largest. There is
a single lateral process on each side near the anterior border of the segment and on
the ventral side there are six processes.

The twelfth segment bears a pair of large spiracles anteriorly, which are placed
in a more dorsal position than those on the other segments. Posteriorly it bears
a row of four large dorsal processes, the median pair being the largest
on the body. On the ventral surface of this segment there is a single pair of
processes.

Fig. 10. Portion of cuticle of Bibio marci larva from dorsal
surface of fourth segment; a, lateral view; 6. surface view ;
the arrows directed anteriorly ; x 330.

The cuticle with which the body is covered bears many small scale-like structures,
which are packed especially closely together on the processes. The largest of these
scales bear stout backwardly-directed spines of a darker colour than the remainder
of the cuticle (fig. 10). The scales on the first, second and third segments, and
some on the ventral surface of the twelfth segment, bear from one to four spines,
and those on the remaining segments usually carry a single stout spine, but a second
more slender spine is present on certain of the scales. The scales on the base of the
processes all carry several spines, but those on the upper part have only a single
spine.

The alimentary canal (fig. 11) takes an almost straight course through the
body, but has a loop near the posterior end, in the hind gut. The largest part of
the alimentary canal is the mesenteron, which bears three large caeca at its anterior
end, one lying on each side and the third, which is the largest, placed ventrally.

THE LARVAL AND PUPAL STAGES OF THE BIBIONIDAE. 227

The four Malpighian tubes join the alimentary canal at the junction of the mesenteron
and hind gut, entering the canal by a short common duct on the dorsal side. There
is no posterior caecum such as is found in connection with the alimentary canal
of Bibio johannis.

Sen yen

re a

Fig. 11. Alimentary
canal of larva of Bibio
marci, lateralview, x 5;

c, caeca; M, Mal-
pighian tubes; oe,
oesophagus.

Pupa.

Elongate, cylindrical, tapering gently to posterior end, abruptly at anterior
end. Completely free from larval skin. Colour at first white, head and thorax of
imago showing dark through cuticle later. Length from head to tip of abdomen,
Bol2-omm: > 9, lo mm. (fig. 12). Head with very slight, blunt, anterior, median
process ; a slight median dorsal ridge extending from the anterior median process
to posterior margin of thorax, dehiscence for emergence of imago occurring along this
ridge. Antennae extremely short, arising between bases of eyes, extending laterally
over eyes. In the males, eyes large and protruding ; in female, smaller and less
conspicuous. Labium elongate, semicircular; labial palpi elongate, conical, extending
laterally. Dorsally, most of head covered by prothorax, only eyes of male visible.

228 HUBERT M. MORRIS.

Thorax strongly arched dorsally. Pair of thoracic spiracles slightly projecting
laterally. Tibiae and tarsi of fore legs extending from side of eyes level with
antennae to anterior margin of second abdominal segment, with tarsi in apposition
along median line. Second and third pairs of legs parallel to first, all except tibiae
and first and last tarsal segments of second and last tarsal segment of third, covered
by wings ; last tarsal segments appearing posterior to those of first leg, and those of
each_pair in contact along median line.

Fig. 12. Pupa of Bibio marct, ds X Se

Abdomen nine-segmented ; length of segments in proportion 7, 10, 11, 12, 12,
11, 10, 9, 9; width of segments decreasing gradually towards posterior end. Cuticle
with many fine wrinkles, mainly transverse; on abdominal segments, dorsally
and ventrally, bearing numerous spines similar to, but less stout than, those of
larva, occurring singly or in groups of two to four, on slight swellings of cuticle.
Terminal segment (fig. 13) bluntly conical, bearing a pair of stout, dark brown,
sharply pointed processes at the posterior end, directed posteriorly and outwardly,
and a pair of rounded papillae. All segments except eighth bearing a pair of slightly
projecting spiracles laterally, towards the anterior margin of the segment.

Fig. 13. Posterior ex-

tremity of pupa of Bibto

marci, 3} dorsal view,
x 18.

Bibio lacteipennis, Ztt.

A number of larvae which proved to be those of Bibio lacteipennis, Ztt., were
found in a plant pot at Withington, Manchester, during February 1918. The larvae
were kept in some of the same soil, which was a rich leaf-mould in which bulbs had
been grown, in the laboratory, and the first pupa was seen on 23rd March, and the
first adult appeared on 13th April.

Larva.

It is proposed to point out the differences between the larva of this species and
that of Bibio marci, L., which has been described above.

The most obvious differences between these two species are the smaller size
of the larva of B. lactetbennis and the fact that the processes on its body are
relatively more slender. The number and arrangement of the processes on all the
segments is the same in both species.

THE LARVAL AND PUPAL STAGES OF THE BIBIONIDAE. 229

The most striking difference is exhibited in the cuticle. In both species the
cuticle of the full-grown larva bears a great number of small, irregularly-shaped,
scale-like structures. In B. marci the largest of these scales bear one stout spine,
while in B. lacteipennis (fig. 14) the scales are much less well developed, and only
those on the processes, and on the anterior part of the ventral side of the first segment
of the body, bear these spines. The latter are, moreover, less stout than those of
B. marci, and there do not seem to be more than two on any scale. On the ventral
surface of the twelfth segment a few of the scales bear single, long, stout spines.

Fig. 14. Portion of cuticle of larva of Bzbio
lacteipennis from dorsal surface of fourth segment :
a, lateral view; b, vertical view; xX 330.

The only apparent difference in the mouth-parts is the presence of a slight median
projection between the two processes at the anterior end of the submentum.

The larva bears a pair of protrusible pseudopod-like structures at the end of the
twelfth segment, which are relatively larger and better developed than is the case
in B. marcv.

The alimentary canal, Malpighian tubes and caeca appear to be similar to those
of B. marci, and there is no posterior caecum.

The length of the full-grown larva is about 15 mm.

Pupa.

The pupa of Bzbio lactetbennis bears a considerable resemblance to that of
B. marci. It is distinguished from the latter by its smaller size and the somewhat
greater development of the median process at the anterior end. The male pupa
bears a pair of small, blunt processes, slightly ventral to the median process and
situated on the anterior portion of the eyes. Length of pupa 7 to 7-5 mm.

Bibio venosus, Mg.
Larva.

A few larvae of this species were found in leaf-mould at Brockenhurst on 22nd
March 1920, but unfortunately before they could be thoroughly examined they
had all pupated.

The mouth-parts do not show any noticeable difference from those of Bibto
marci, and the number and arrangement of the spiracles and processes is similar.
With this larva again, as in the case of B. lacteipennis, the chief difference from
B. marci is in the scales on the cuticle. As in B. lacteipennis, these scales are very
slightly developed and do not bear spines, except those on the twelfth segment

230 HUBERT M. MORRIS.

and occasional ones on other segments, which bear a single, short, blunt spine. The
scales in this larva are much more regular and are placed more closely together than
in any other species so far examined (fig. 15).

=) 6

Fig. 15. Portion of cuticle of larva of

Bibio venosus from dorsalsurface of fourth

segment: a, lateral view; b, vertical view ;
<7330:

Pupa.

This pupa again bears a considerable resemblance to that of B. marci, from which
it also differs in its smaller size; in the greater development of the process at the
anterior end, which is slightly larger than that of B. lactetpennis; and in the
presence in the male of a pair of similar blunt processes on the anterior portion
of the eyes. The eyes in the male of this species are rather smaller than in the
two species previously described. Length of pupa 10 to 11-5 mm. In the
laboratory the pupal state lasted 19 days.

Differences between Larvae of various Species.

In the foregoing descriptions of Bibio larvae it will be noticed that attention
is drawn to the differences in the cuticular structures in the different species dealt
with. These differences seem so far to be sufficiently distinct and regular to give

Fig. 16. Portion of cuticle of larvaof Bibio johannis from
dorsal surface of fourth segment: a, lateral view;
b, vertical view; x 330.

a ready means of distinguishing between the species in the larval state. There
are many other species of bibio, the larvae of which have not yet been examined,
but it is possible that some such distinction may hold good throughout.

THE LARVAL AND PUPAL STAGES OF THE BIBIONIDAE. 231

In addition to the three species described above, larvae of Bbzbio johannis,. L.
(fig. 16), have been examined, and also larvae believed to be those of B. pomonae, F.
(fig. 17), but the latter were not definitely identified in the adult state. As will be
seen from the illustrations, these two species are quite distinct from the other species
figured,

Fig. 17. Portion of cuticle of larva of Bibio pomonae from
dorsal surface of fourth segment: a, lateral view;
b, vertical view ; the arrows directed anteriorly; x 330.

The alimentary canal also varies in some of the species examined, that of
B. marci and B. lacteipennis being without the posterior caecum which is present in
the canal of B. johannis. This posterior caecum is also absent, according to
Keilin (4), from the alimentary canal of B. hortulanus, L. The labium also shows
slight variations in different species.

Parasites of Bibionid Larvae.

Very few parasites of the BrpiontDaE are known. In addition to the mite
and nematode recorded by Lyonet, Keilin (4) mentions the Gregarine Schneideria
mucronata, Leger, occurring in the mid-gut and anterior caeca of Bibionid larvae,
and also a bacterial disease. He further mentions a Microsporidian, probably a
Glugea, which invades the epithelial cells of the mid-gut and caeca of Scatopsid
larvae.

Sorauer (7) gives Agyrtes bicolor as a parasite of the larvae of Scatopse.

A Gregarine, probably the Schneideria mentioned by Keilin, was found in the
posterior caecum of a Dilophus febrilis larva, and in the same larva two cysts full
of spores were found free in the body. Another parasite, probably the Glugea
mentioned above, was found infesting the epithelium of the alimentary canal in a
Bibio johannis larva.

In addition to these a Dipterous parasite was met with in connection with
Bibio marci larvae. This was the Phorid Hypocera incrassata, Mg. The fully grown
larvae emerged from the B. marci larvae when the latter were about to pupate,
and almost immediately entered the pupal state. Only a single parasite was observed
in each larva attacked.

Additions to Literature since 1917.

In 1917 an account of the larva and pupa of Bibio johannis was published, in
which the literature up to that date was referred to. The following publications
have appeared since that date.

232 HUBERT M. MORRIS.

A brief account, with figures of the larva, pupa and adult, of Plecia fulvicollis, F.,
an Indian Bibionid, has been published (3). Keilin (4) has given a short account
of the alimentary canal in Scatopse notata, Bibio hortulanus and Dilophus febrilis.
That of B. hortulanus resembles the alimentary canal of B. marci in the absence of
the posterior caecum, which however is present in a well developed state in Dilophus
febrilis.

A few additional records of damage to crops by Bibionid larvae have been published
also. Carpenter (1) records Bibio larvae as having caused much damage in wheat
fields in Co. Cavan, Ireland, in March 1918, by eating the roots of the plants; and
Bibio marci larvae are reported to have damaged potatoes in a “ pit” in Co. Louth,
in October 1916.

Molz (5) records Bibio hortulanus larvae as damaging potatoes near Halle in
September 1918 by eating beneath the skin. It was noticed that only parts of the
field which had been treated with stable manure were infested. Afterwards winter
wheat was sown and larvae were found to be feeding on the seed. In experiments it
was found that the larvae attacked the seed at the point of germination, and also that
only potatoes which had previously been injured were attacked. Potato skins
steeped in a 1 per cent. solution of arsenious acid were found to form an attractive
and effective poison bait.

1. CARPENTER, G. H. (1920). ‘‘ Injurious Insects, etc., in Ireland during 1916,
1917 and 1918.” Econ. Proc. Roy. Dub. Soc., ii, No. 15, Nov. 1920.

2. Curtis, J. (1844). ‘‘ Bibio marci.” Gardeners’ Chronicle, 16th Nov. 1844,
p. 764.

3. FLetcuer, T. B. (1919). ‘‘ Second Hundred Notes on Indian Insects.”” Agric.
Research Inst., Pusa, Bull. 89, p. 58.

4. Kein, D. (1919). ‘‘On the Alimentary Canal and its Appendages in the
Larvae of Scatopsidae and Bibionidae (Diptera Nematocera), with some Remarks
on the Parasites of these Larvae.’’ Ent. Monthly Mag. (3) v, pp. 92-96.

5. Morz, E. (1920). ‘‘ Weitere Beitrage zur Kenntnis der Gartenhaarmiicke (Bzbi10
hortulanus, L.).”’ Zeitschr. angew Entom. Berlin, vii, No. 1, Sept. 1920. pp. 92-
96, 3 figs. (Abstract in Rev. App. Ent., 1921, p. 1).

6. Morris, H. M. (1917). ‘The larval and pupal stages of Bibio johannis, L.”
Ann. App. Biol. iv, 3, pp. 91-114.

7. SORAUER, P. (1913). Handbuch der Pflanzenkrankheiten, iii, p. 458.

Butt. ENT. RESEARCH. VoL XII. Part 3. PEATE VT:

Fig. 1. A typical ease of ‘‘ Simulium disease.”’

Fig. 2. Chemosit River flowing through forest heavily
infested with Simulium.

BuLt. ENT. RESEARCH. VoL. XII. Part 3. PLATE VIII.

Fig. 1. Dense thorn forest heavily infested with Simulium, Lumbwa
Reserve, Kenya Colony.

Fig. 2. Thin thorn forest, adjoining that in Fig. 1, lightly infested
with Simulium.

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233

TRYPANOSOMIASIS IN THE ABSENCE OF TSETSES, AND A HUMAN
DISEASE POSSIBLY CARRIED BY SIMULIUM IN KENYA COLONY.

By F. W. Dry, M.Sc.,
Recently Asststant Government Entomologist, Kenya Colony.

(Prates VII -& VII).

This is an account of two different inquiries carried out on the same safari in the
Lumbwa country of Kenya Colony early in 1920. The problems are of the same type,
the suspected transmission of disease by blood-sucking flies, and the second investiga-
tion arose out of the first. It therefore seems best to deal with them both in a single
paper.

In March 1920, in the absence of my Chief, Mr. T. J. Anderson, I was asked to make
a fly survey in the neighbourhood of a farm in the Kericho district where cases of
trypanosomiasis had been demonstrated. I arrived at this farm on 25th March. The
blood-sucking flies collected are listed below. As was anticipated, the altitude being
about 6,300 ft., and there being no records of tsetse-fly anywhere near, none
was found. I was then fortunate in receiving information from Mr. Ian Q.
Orchardson, the pioneer settler of the Kericho district, about a small blood-sucking
fly to be found in the forest in part of the Lumbwa Reserve. The bites of this fly,
he told me, are believed by the Lumbwa to be the cause of a disease from which some
of the inhabitants of that region suffer. He further suspected that this fly might
transmit a disease of cattle. Accordingly, on 7th April, I proceeded into the area
about which he had told me in the Lumbwa Reserve, also crossing the Kipsoni into
adjacent country of the Kisii Reserve. The fly proved to be a Simulium. I satisfied
myself that it could not be regarded as the transmitting agent of any cattle disease,
but did obtain evidence which shows that the belief of the Lumbwa calls for further
investigation. On 29th April, I returned to the farm near Kericho, to search for
Simulium there, but failed to find it, staying there until 3rd May.

I. Possible Transmission of Trypanosomiasis in the Absence of Tsetses.

The presence of trypanosomiasis on the farm near Kericho was demonstrated
from blood-slides taken by the late Captain J. Cameron, Veterinary Officer, and
examined by Mr. W. Kearney, Assistant Veterinary Pathologist. His diagnosis
of the type of trypanosome was confirmed by Mr. R. E. Montgomery, now Veterinary
Advisor in Uganda, who wrote to Mr. Kearney in January 1920 :—‘ On purely mor-
phological features, the trypanosomes... .. are recognisable as Ty. untforme.
The bold body, slightly developed, undulating membrane, short, free flagellum, dis-
tinct blepharoplast and clear cytoplasm all point to this species, or a near relative
of the vivax group.”

The farm where the outbreak occurred was a new one, and all the cattle on the
farm, other than those of native squatters, were working oxen which had been bought
and brought on to the farm towards the end of 1919 or early in 1920.

In 1919 there had been no deaths of oxen, apart from one animal which died
after an accident, but from early January up to my arrival there had been a high
mortality, and trypanosomiasis had been demonstrated from blood slides in some
cases. Particulars about the losses up to the time of my visit are as follows :—

Total number of European owned oxen - Ae in 63
Deaths :—
Trypanosomiasis proved by blood-slides 2
Not due to any disease .. ae ; 2
Cause not proved .. ; 14

234 : F, W. DRY.

Ten cases of trypanosomiasis were proved from blood-slides, all taken previous
to the 6th April, and eight of the oxen were still alive at the time of my visit.

After my visit the mortality ceased. No more cases of the disease were detected,
and all animals proved to be infected were slaughtered in order to bring the outbreak
to a close.

It should here be mentioned that two farmers resident for some years in the district
told me that when cattle are brought from other parts of the country to poor grazing
of the sort on this farm and worked hard, as the oxen had been, there is great risk of loss.
This, it seemed, might be the explanation of some of the deaths. With the fact,
however, that, in all, ten cases of trypanosomiasis were proved, and that no other
disease had been demonstrated on the farm, the presumption certainly is that some
of the fourteen deaths of which the cause was not known were due to trypanosomiasis.

There were no signs of the disease amongst the small number of cattle owned by
Lumbwa on the farm. The European and native owned animals were kept away from
each other.

The European owned oxen had all been bought from dealers, two European
and two Indian. Those obtained from the European dealers had been through the
Kibigori Quarantine Station, to which animals were taken from widely separated
sources ; those from Indians had probably come out of the Lumbwa Reserve.

Of the ten animals proved to have trypanosomiasis two had been purchased
from one European dealer, two from the other, and the remainder from one or
other of the two Indian dealers. <A sufficiently accurate record had not been kept
to make it certain whether animals which had the disease had come from one
Indian dealer or from both, but the partner on the farm in charge of the cattle felt
almost certain that it was from both. At any rate, animals from three out of the
four sources had trypanosomiasis, and though there is just a chance that both the
European dealers had bought cattle from the same mob from Kibigori, the number
of distinct sources from which animals proved to have trypanosomiasis had come
is at least two, though it may be four.

Another point to be noted is that the second animal which was proved to have the
disease came on to the farm early in February, having been purchased from one
European dealer, after the death, on Ist February, of the first animal proved to have
the disease, which had been bought from the other European dealer.

One piece of information given to me must now just be mentioned about the
possible presence of trypanosomiasis cases other than oxen on the farm. A Dutch
manager who had left the farm had had two donkeys. One donkey is believed to
have died or been killed on the farm in November or December of 1919. The other
had been given to a Kericho resident and had subsequently died. This manager
was stated to have thought that the donkeys had trypanosomiasis. This
evidence is not of a definite description, but is recorded because Captain Cameron
inclined to the view that it might have been in these donkeys that the disease was
brought on to the farm.

I collected blood-sucking flies on this farm and on neighbouring land, near to the
river known at different parts of its course on the farm boundary as the Saosa, the
Dimbilitch, and the Kitho, near to small tributary streams, and on the grazing
grounds of the cattle, which were then being kept away from the river. The
blood-sucking fly fauna could not be considered a striking one. All such flies
captured belonged to the genera Haematopota and Stomoxys. Stomoxys were chiefly
captured on grazing oxen, often being present in large numbers, while almost all the
Haematopota were taken near to the river.

TRYPANOSOMIASIS IN THE ABSENCE OF TSETSES. Zao

While in the Lumbwa and Kisii Reserves I was collecting blood-sucking flies
at various lower points on the same river system, as far as a point on the Sondo
River about fifteen miles in a direct line from the farm, and with the exception of a
Simulium, to be referred to shortly, the only specimen taken of a fly of any
additional genus was a single individual of Tabanus taeniola, P. de B., var. variatus,
Walk., captured in the Lumbwa Reserve about ten miles due west of the farm.
While it is not impossible that that fly might be taken on the farm, it can hardly
be common there. The Lumbwa herd-boys on the farm told me that my captures
included specimens of all the blood-sucking flies which they knew to be there.

The cattle on the farm were not dipped and some ticks were collected from them.
The species of blood-sucking flies and ticks taken were the following :—

Haematopota similis, Ric—10 929.

; hirta, Ric.—2 @ 9.
brunnescens, Ric.—11 9°.
alluaudt, Surc.—5 @ 8.

* ugandae, Ric.—28 92 9.
Stomoxys calcitrans, L.—15 $4, 22 29.

varipes, Bezzi—3 92 9.
= nigra, Macq.—1 Q.
Amblyomma variegatum, F.
Boophilus sp.

Mr. Orchardson told me of an extensive area of thick thorn forest in the Chemosit
River district of the Lumbwa Reserve which the Lumbwa leave almost uninhabited
owing to its being infested with a fly which proved to be a Simulium. The belief of
the Lumbwa, about which he told me, that the bites of this fly cause a disease of
human beings will be discussed directly. He further suspected that this fly might
be the carrier of some disease of cattle. In 1917 he and a neighbour carted maize
to their farms near Kericho from Litun, and outspanned their oxen just off the Sotik
Road near to the Chemosit and Jamji Rivers. Afterwards some of the oxen used
in this work died, each planter losing about half a dozen animals. “‘ These,” he
said, “ gradually faded away and died.” Some were ill for weeks, some for months.
They were not seen by any veterinary officer. Now there is a considerable area
of thorn forest on and near the farm where the outbreak of trypanosomiasis had
occurred, so this was a suggestion to be followed up.

My inquiry, however, yielded negative results, for :—

(a) Those Lumbwa who live in the infested area keep cattle. One young man
had come two years ago from a district clear of Simulium in order to
graze his father’s cattle in the infested country, for much of the grazing
is good.

(b) The people told me that Simualium bites cattle, sheep and goats, but they
said that it does not give the animals any disease.

(c) It was pointed out to me by Mr. C. M. Dobbs, the District Commissioner
at Kericho, that cattle are frequently outspanned just off the Sotik Road
in the same place as those of Mr. Orchardson and his neighbour, but he
had not heard of any other cases of disease supposed to have been
contracted there like that of which Mr. Orchardson had spoken.

(d) I failed to find Simulium so far east as the Sotik Road, the nearest point
where a specimen was captured, and that at a place where Simulium
was not numerous, being a mile and a half to the west, though it is not
impossible that the fly may occur on the road in small numbers.

After this visit to the Reserves I returned to the farm of the outbreak and again
collected blood-sucking flies there and in the thick thorn forest on adjacent land,

236 F. W. DRY.

searching especially for Simulium but finding none. Inquiries were made from a
large number of natives, who replied, either that they knew nothing at all about
the fly, or that they knew of it near the Chemosit River.

Simulium was therefore not a factor in the presence or transmission of trypano-
somiasis on the farm where this outbreak had occurred.

The evidence recorded above, combined with the fact that no other outbreak
of trypanosomiasis was recorded in the district, points to the probability that the
disease was transmitted on the farm to healthy animals from one or more animals
which were infected when they came on to the farm. If that be so, suspicion would
fall first on Stomoxys.

Il. Possible Connection of Simulium with a Disease of Human Beings.

The Lumbwa name for this Simulium is “ kekonjek.” A place infested with
Simulium is spoken of as ‘‘ kapkekonjek.’”’ The insect has been identified by
Mr. F. W. Edwards, of the British Museum, as Simulium neavei, Roubaud.

The badly infested country, so far as I was able to ascertain from my own
observations and from inquiries from the Lumbwa, is covered with dense forest,
mostly of thorn trees, white and red, with occasional open spaces (see Plates vii
and viii). It is well watered, and it is unlikely that there is any point within that
area much more than two miles from some stream, large or small. About three
miles below the junction of the Chemosit and Kipsoni Rivers one fly was captured,
this being the lowest point reached on that river on the trip ; Szmuliuwm was present
there in only small numbers and it was evident that, lower down, neither the Lumbwa
on the right bank of the river, nor the Kisii on the left, paid any attention to the
fly. It was also taken in small numbers in a tributary valley of the Sondo River
on the left bank a mile from the Sondo ; in this valley there were scattered clumps
of trees but not continuous forest.

The flies are appreciably less numerous in thin than in thick forest (see Plate viii),
and while present in the middle of open spaces—say a quarter of a mile in diameter—
are there distinctly less numerous than in the forest. This falling off in numbers as
one gets away from dense forest is quite a sudden one.

The flies are active, the Lumbwa told me, and my own observations are consistent
with what they said, from about seven o’clock in the morning until five in the after-
noon, and especially so in the afternoon. Very little is seen of them when rain is
falling, and I was told, and from my own collecting believe it to be so, that the flies
are not so active in intense sunshine as when the sun is less bright. They pay little
attention to people on the march, but if one remains still in infested country they
appear immediately. They bite very readily and often raise half-inch wheals, a
drop of blood frequently exuding from the puncture. In badly-infested forest a
person sitting with a bare back and allowing the flies to do as they liked would
probably be bitten once a minute at the very least.

It is the accepted belief of the Lumbwa and Kisii amongst whom I moved, that a
disease is caused by the bites of these flies, the chief symptom being that the skin is
in folds (Plate vii, fig. 1). It would seem that these folds appear first on the small
of the back, but sometimes the skin higher up the back, and more rarely that of the
front and sides of the body was seen to be affected in the same way, while occasionally,
though this was only so with elderly people, the condition would extend to the arms
and legs. Sometimes the skin was spotty, as shown in the photograph, and occasional
cases were shown to me as Simulium disease in which spots, but no folding, were
present. People with the affection scratch themselves a great deal.

Many of the affected people that I saw were able to carry on their usual activities,
but some looked to be in quite a feeble condition. I asked one young man if he

TRYPANOSOMIASIS IN THE ABSENCE OF TSETSES. 237

could work properly, and those present laughed at my putting such a question to him.
One man, the one shown in the photograph, I spoke of as “ elder,” and was told that he
looked like an elder because of the disease, but was really only a “ warrior.”

Poor sight, or even blindness, is stated by the Lumbwa to be caused by the
disease occasionally, but two people with the skin affection who had bad sight looked
to me to be suffering from cataract.

The youngest person with the skin affection that I saw was a boy perhaps sixteen
years old, but I was told that quite small children sometimes had it. Young people
may recover from the disease, and I met three people who said they had done so ;
one of them, now an elder, said he had been ill for five years, and recovered ten years
ago. With old people, I gathered, the condition always persists until death. Some
people told me that they had had the disease for thirty years or more. It seems
unlikely that the disease is the cause of death.

While the Lumbwa are convinced that this disease is caused by the bites of Simu-
lium, they do not believe that it can be caused by a single bite or by only a few bites.
I was told repeatedly that a person living a short time, say a fortnight, in badly
infested country, would not get the disease, and people that I met have lived there
much longer than that without contracting it. Again, I was told that while the
disease could be contracted by herding animals in infested country day after day,
it would be safe for a person to herd them there occasionally. The Lumbwa have no
hesitation in going into infested country now and again to hunt, pick vegetables, or
search for honey, and a very well-marked track runs right through the forest between
the well-inhabited districts to the north and to the south of the infested forest on
both sides of the Chemosit. And further, just away from the edge of infested forest,
and in country where I took Simulium in small numbers, many Lumbwa were living,
and considered themselves to be running only very slight risk of getting the disease.
Simulium, the Lumbwa told me, does not enter houses.

Part of this slightly infested area adjacent to badly infested forest, the Lumbwa
told me, they had cleared of thick forest, thus, they said, themselves converting it
from unsafe into reasonably safe country. Another obvious way of avoiding the
bites of the fly is the wearing of European clothes. I was scarcely bitten at all
myself while moving in infested country for three weeks.

The total number of people that I saw who were said to have the disease at the
time of my visit, not counting one Kisii, who possibly had the disease and who will be
mentioned again shortly, was twenty-three. Seven other cases I heard of, but did
not see, and probably some others exist. Of these twenty-three people, twelve were
male and eleven female. Roughly speaking, eleven were young and twelve old. Of
the twenty-three, eight were living in the forest at the time of my visit and at least
six had lived there previously, making their homes elsewhere after getting the disease.
The remainder had lived near to infested forest and had often gone into the forest.
I was told that cases of this disease were not to be found in country away from the
vicinity of infested forest. It was said that if a person with the disease attempted
to settle in country other than adjacent to the infested forest, he would not be allowed
to do so by the other inhabitants.

The one apparent exception which I came across, to the coincidence of distribution
of the disease and Simulium, was a Kisii elder whose skin showed both folds and spots,
but who was stated never to have lived in infested country.

Special reference must be made to two settlements in clear spaces in the infested
forest, areas in which the fly could be found in fair numbers, and both surrounded
by heavily infested forest, where I inquired, as far as I was able, after the health of
those who were living there. There may have been other people that I did not see
or hear about, but eight adults that I saw who had lived there for more than two years
all had the disease, though one other old resident, a man that I did not succeed in

(4183) S

238 F. W. DRY.

seeing, was stated probably not to have the disease. Six other adults who had lived
there from a few weeks to two years were healthy. A number of children, too, some
of whom had lived there all their lives, and might be as old as ten, were all free from
the disease ; they do not go into the forest nearly as often as grown-up people.

These facts, therefore, do indicate coincidence of distribution of the disease and
Simulium.

One piece of tribal history that I learnt from both the Lumbwa and Kisii throws
light on the way in which Simulium is regarded by these two peoples. Formerly
the country badly infested with Szmuulium, which is now in the Lumbwa Reserve, was
occupied in its western part by the Kisii, the eastern part being left uninhabited out
of mutual fear, for in those days the relations between the two tribes were by no means
peaceful. Some time before the area came under British influence, possibly, though
such information given by natives must be considered of doubtful accuracy, about
thirty years prior to the present day, the Lumbwa succeeded in driving out the
Kisii from the infested country, where they proceeded to settle, and established
themselves, too, on the left bank of the Kipsoni and Sondo, but these Lumbwa were
moved back across the river by the British Government.

I had a lengthy conversation with some of the Kisii, now old men, but warriors
at the time of the war, who had formerly lived in the infested country. They told
me that they knew the fly and the disease and believed the one to be the cause of the
other. The Kisii had not hesitated to live in that country on account of the disease.
Only a few of their people, they said, had had the disease. The land, they said, and
the Lumbwa had told me the same, was excellent for crops and the grazing good.
Wishing to know how seriously they regarded the disease, I asked them if they would
go back to the Simulium country if they could. They replied, ‘‘ Certainly,” and asked
if I had been sent to arrange for their return.

After that war the Lumbwa settled in the conquered country in considerable
numbers, but very soon found they were getting the disease, when the great majority
left the district. A few of the Lumbwa living in infested country and having the
disease were amongst the original settlers and a few more have settled there more
recently, but the total number of inhabitants, in an area of perhaps 20,000 acres, is
small in the extreme.

I obtained blood-slides from three Lumbwa men who had lived in the infested
country ever since it had become Lumbwaterritory, and said they had had the disease
for about thirty years. These slides Dr. Clearkin, Acting Senior Bacteriologist,
afterwards kindly examined, finding about 20 per cent. of eosinophilia in the blood
from each of the three individuals.

Dr. Clearkin hopes to find an opportunity of visiting the Lumbwa country to
investigate the disease. The District Commissioner has tried to persuade some of the
Lumbwa having the disease to come to Nairobi in order to be examined, but they
have declined to leave their Reserve.

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SPIRACLES OF LARVAE OF MYIASIS-PRODUCING FLIES.

239

NOTES ON THE MYIASIS-PRODUCING DIPTERA OF MAN AND
ANIMALS.

By Major W. S. Patton, M.B., F.E.S., I.M.S. (rtd.),
Lecturer on Entomology and Parasitology, Edinburgh University.

(PLATES V & VI.)

Myiasis has been defined by Austen as follows :—‘‘ There remains yet a third
category of flies, chiefly belonging to the great Family Muscidae, the larval stage of
which is sometimes actually passed in the living human body, the presence of the
larvae in the various organs and tissues, and the disorders or destruction of tissue
caused thereby, being comprehensively known as Myiasis.”’ It will be noted that this
definition only refers to the larval stages of flies, but as the eggs, and in one case at
least, pupae and adults, may be found in the human body, and as myiasis is a common
condition in many animals, the definition may now be amplified as follows :—The
condition or conditions, resulting from the invasion of the tissues and organs of man
and other animals by all the stages of the Diptera.

In the latest edition of Castellani and Chalmer’s ‘‘ Manual of Tropical Medicine”’
(1919) species of flies belonging to no less than ten families have been incriminated
as Causing myiasis in man ; there is very little doubt, however, that the identification
of some of the species is more than doubtful. These lists do not even include the
most important myiasis-producing Calliphorine of Africa, Chrysomyta bezziana, Vill.,
so that the information given on this subject must be considered very incomplete.
This is chiefly due to the fact that it is difficult to breed the flies from larvae recovered
from cases of myiasis, in order to be sure of the identity of the species, and unfortu-
nately in most cases identification of species by larvae alone is not yet possible.
Although specimens of larvae are doubtless plentifulin museums and private collections,
their respective adults, accurately identified, are, on the other hand, scarce.

Quite recently I undertook an inquiry into the larvae of the myiasis-producing
Diptera of India in order to discover to what species of flies they belonged. Through
the kindness of the Indian Research Fund Association I was able to issue a printed
appeal, with detailed instructions for collecting living larvae, to every medical and
veterinary officer in India, Burma, Assam and Ceylon, and in a very short time large
numbers of living larvae and puparia were received from all over this area, and hun-
dreds of flies were bred out and accurately identified. The results of this inquiry
have demonstrated how little we know of the Indian species whose larvae cause
myiasis in man and animals. The following species were found to be those which
usually cause myiasis in man and animals in India :—

(1). Chrysomyia (Pycnosoma) bezziana, Vill—This proves to be the common
myiasis-producing Calliphorine of India and adjacent parts. Its larvae cause rhinal,

oral, ocular, aural, vaginal and cutaneous myiasis in man and animals, but its larvae
never cause intestinal myiasis.

(2). Chrysomyia megacephala, Fabr. (dux, Esch.) and Lucilia argyricephala, Fabr.
(serenissima, Macq).—The larvae of both these species may be occasionally found
in cases of myiasis in animals.

(3). Sarcophaga sp. incert.—This species, which it has not: been possible to identify,
as the larvae were sent to me preserved in spirit, occasionally causes cutaneous and
intestinal myiasis in man. In the former cases the destruction of the tissues is very
extensive.

(4183) S2

240 MAJOR W. S. PATTON.

(4). Aphiochaeta xanthina, Speiser (ferruginea, Brunetti), and A. rufipes, Meig.—
These two Phorids commonly cause tissue and intestinal myiasis in man and animals
in their larval stages.

During the course of this inquiry, which extended over more than a year, I gained
a large amount of valuable information on such points as, how best to breed these
flies from their larvae, how to preserve and send the larvae for further study and, most
important of all, how to set about identifying the larvae and adults of blowflies in
general, and the myiasis-producing species in particular. It is with the object of
extending the inquiry to the study of the myiasis-producing flies from other parts of
the world, particularly from Malaya, the Dutch East Indies, China, Japan, Australia
and Tropical America, that I propose recording my notes in this paper. Although
much valuable work has been done on the myiasis-producing Diptera of Africa by
Rodhain, Bequaert, Roubaud and others, our information regarding the CaLtt-
PHORINAE which parasitise man in their larval stages is still very imperfect. For
instance, it is not known whether Chrysomyia bezziana deposits its eggs in human
tissues in Africa, though its larvae are extremely common in human tissues in India.
I would therefore ask all those who may have opportunities of collecting such material
from man and animals to forward it to Dr. Guy A. K. Marshall, C.M.G., Director
of the Imperial Bureau of Entomology, British Museum (Natural History), Cromwell
Road, London, S.W. 7., who has kindly consented to receive it and forward it to me
for further study. The results will be published from time to time in this Bulletin.
I may say that as the identity and correct names of many of the common blowflies
are at present hopelessly confused, I have decided to extend this study to the CaL-
LIPHORINAE in general. Specimens of any blowflies and their larvae, particularly
first stage larvae, and any of the larvae and adults of the OESTRIDAE will be most
welcome. Most of the Indian material has already been collected, and is now being
studied ; material from the areas mentioned above would be of great value for
comparative study. Those who wish to help in this inquiry will find full details of
how to collect and breed blowflies in general, and the myiasis-producing species in
particular, in these notes. I shall be glad to correspond with any observer who would
like any further information on this subject.

The Classification of Myiasis-producing Diptera.

A reference to the recent literature on this subject shows that the myiasis-pro-
ducing Diptera are mainly grouped according to the tissues or organs in which their
larvae are found, so that we have such species as cause rhinal, oral, aural, ocular,
tissue, and cavity myiasis. But such a Calliphorine as Chrysomyia bezziana will
deposit its eggs in any of these organs or tissues, so that following this method of
classification it would have to be placed in several of the groups.

Recently Bishopp has contributed a valuable article on this subject, grouping
the species according to the methods of attack , his groups are as follows :—

(1). Tzssue-destroying forms.—In this group are included those species whose
larvae attack living animals secondarily, mainly breeding in the bodies of dead animals.
Bishopp rightly places Cochliomyia (Chrysomyia) macellaria, the well-known screw-
worm fly of Tropical America in this group, also such species as Lucilia sericata, and
species of Sarcophaga. Chrysomyia bezziana would also have to be placed in this
group, for its larvae are true tissue-destroyers, as anyone who has seen a scalp wound
infested with its larvae will know. But this Calliphorine cannot be placed in the
same group as Cochliomyia macellaria, Lucilia sericata and species of Sarcophaga,
for the simple reason that it never breeds in dead tissues ; it is a true specific myiasis-
producing species, and the only known metallic Calliphorine which has this habit.

(2). Subdermal migratory forms.—Species, the larvae of which are true parasites
of man and animals during the greater part of their lives, living beneath the skin,

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 241

and in the subcutaneous tissues; Dermatobia hominis, Hypoderma bovis and
Cordylobia anthropophaga are good examples. Chrysomyia bezziana must also be
placed in this group, for its larvae are true dermal parasites of man. I have had
examples of its larvae sent me from such small skin punctures as are made in boring
the lobes of the ears for ear-rings, and in small cuts on the skin. The only difference
between a case of myiasis caused by the larvae of Chrysomyta bezziana and that of
Cordylobia anthropophaga is that, in the former, it is a massed infection, while in
that of the latter it is only one larva which causes the myiasis. In the former the
tissues rapidly break down and suppurate, while in the latter they do not. Both
are specific myiasis-producing Diptera.

(3). Larvae infesting the intestinal and urogenital tracts——The larvae of these
species feed, to a lesser or greater extent, on the food contained in these organs,
especially in the gastro-intestinal tract. In this group Bishopp includes those species
whose larvae accidentally find their way into the intestinal tract, such as the larvae of
Fannia canicularis and those of many species of Sarcophaga, and those OESTRIDAE
whose larvae live in the intestinal tract of animals. I should like to point out that
two categories are here involved ; in the one case, the larvae only accidentally pass
into the gastro-intestinal tract, the species concerned normally breeding outside the
human body, whereas in the case of the OESTRIDAE mentioned, the larvae can oly
live in special parts of the alimentary tract. Species with such distinct habits cannot
be placed in the same group.

(4). Forms infesting the head passages.—In this group Bishopp includes those
true parasites of man and animals, the larvae of which live in the nose and its acces-
sory sinuses, the throat, etc. Here again, this group would have to include Chrysom-
yia bezziana, for its larvae also live in the nose and accessory sinuses, and are true
human and animal parasites. In the one case the tissue destruction is not great,
Whereas in the other it is often extensive.

(5). Blood-sucking forms.—In this group Bishopp includes those species whose
larvae are blood-suckers. Blood-sucking can, however, hardly be considered a form
of myiasis, for then we would have to include in this group all the blood-sucking
Diptera.

I have said enough to show that the various methods of grouping the myiasis-
producing Diptera according to the tissues or organs attacked, or the methods of
attack, are not satisfactory. I am of opinion that the subject is best considered
from the standpoint of the habits of the flies themselves, which then naturally fall
into three groups as follows :—

(1). Specific myiasis-producing Diptera.

(2). Semi-specific myiasis-producing Diptera.

(3). Accidental myiasis-producing Diptera.

I will now give a few notes on the adults, hosts, geographical distribution, etc.,

of the species belonging to the various groups, reserving the notes on all the larvae
until their identification is dealt with.

Specific Myiasis-producing Diptera.

In this group I include all those Diptera whose larvae are found only in living
tissues, the flies selecting a number of tissues or organs, or one particular organ,
depending on the species, in which or near which to lay their eggs or deposit their
larvae. The following species are included in this group :—Chrysomyia bezziana,
Cordvlobia anthropophaga, C. rodhaini, Wohlfahrtia magnifica, and all the OFSTRIDAE.

Chrysomyia bezziana, Villeneuve.-—As I have mentioned above, this species
is the specific myiasis-producing Calliphorine of India, Burma, Assam, Ceylon and
Africa. Its larvae are commonly found in the nose and accessory sinuses, the mouth,

242 MAJOR W. S. PATTON.

ear, eye and vagina, and in sores, cuts, wounds and abscesses, in man and animals.
As a result of the enquiry made in India, I received the larvae of this species from
more than 170 cases of human and animal myiasis; not a single larva was sent
from the intestinal tract. As this fly breeds only in living tissues it is rarely seen in
nature, for it does not frequent places where most other blowflies may be seen,
such as food stalls, refuse heaps, latrines, and about the bodies of dead animals.
The female fly is attracted by any offensive discharge, and will then lay her eggs
in a mass, varying in number from 380 to 500, on or near the tissues from which the
discharge comes. A sore, wound, or a diseased organ, such as a nose from which
there is an offensive discharge, is therefore a necessity. The larvae rapidly reach
maturity, owing to the high body temperature and rich food, and when mature
crawl out and pupate in the ground. It is very probable that the adults are flower-
feeders, and that as the mature larva stores up a large amount of food in its fat-
body, there is sufficient nutriment for the eggs to ripen, as in the case of the OESTRIDAE.

Chrysomyia bezziana is a well-known pest in Africa, where it has been recorded
by Rovere, Bouet, Roubaud and Joyeux from the Belgian Congo, the Upper Ivory
Coast and French Upper Guinea; by Jack from Rhodesia; and by Aders from
Zanzibar. Curiously enough, in Africa its larvae have been recorded only from the
tissues of the larger animals, chiefly cattle ; so far as I am aware, they have never
been found in human tissues.

In India the female C. bezziana may be readily confused with the female of
Chrysomyia megacephala, F. (dux, Esch.) ; both species have yellow cheeks. In
order to distinguish them with any certainty, it is necessary to examine’ the front
and to note the following points :—The front of the female megacephala is more than
one-third the width of the whole head, that of bezzzana is distinctly less than one-
third. The frontal stripe of megacephala is wider than that of bezziana, the sides
appearing slightly convex along their outer margins ; in bezziana the stripe is slightly
browner and the sides are almost straight. The para-frontals in megacephala are
wider and of a greyish yellow colour, those of bezziana being much more silvery.
The ocular margin of megacephala is slightly concave, and the lower margin of the eye
rounder than in bezziana. The vertical bristles of megacephala are better developed
than those of bezziana. Lastly, and this is perhaps the most valuable distinguishing
character, the squamae of megacephala are of a dirty yellow colour, while those of
bezziana are waxy white; in both they are covered with dark hairs.

The male megacephala can be readily distinguished from the male bezzzana by its
bright red eyes, and the area of large lenses occupying about the upper third and
surrounded by smaller ones. In the male bezziana the eyes are brown, and the
lenses are small and all of about the same size.

Although originally described from Guinea, I have not seen any specimens of
megacephala from any part of Africa, so I am unable to say whether it occurs there.
It is plentiful in the bazaar at Port Said, and one would expect it to occur in Egypt
at least. It is, however, present in the Australian region.

There is no other Calliphorine which may be confused with bezziana, unless it be
Chrysomyia flaviceps. I have not been able to identify this species with certainty,
and whenever this name is used by writers in India, Chrysomyia megacephala is
meant, so that there is some confusion regarding it; it is probably a synonym
for megacephala.

There is much which has yet to be discovered in connection with the distribution
and life-history of Chrysomyia bezziana. For instance, we do not know the limits
of its eastern distribution. Does it occur in Malaya, the Dutch East Indies, China
and Japan? I hope that those who have opportunities of collecting larvae from
living tissues, either of man or animals, in any of the areas mentioned above, will
do so, and send me the larvae for identification,

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 243

Cordylobia anthropophaga, Blanchard.—This is the most important specific
myiasis-producing Calliphorine of Africa, where it is widely distributed, ranging
from the Bahr-el-Ghazal Province of the Anglo-Egyptian Sudan in the east, and the
northern limits of Senegal in the west, to Natal in the south. The most complete
account of its life-history and habits will be found in Roubaud’s, “ Etudes sur la
Faune parasitaire de l'Afrique Occidentale Frangaise,” Part 1. In its larval stage
it is essentially a parasite of dogs, but is also found in a number of other domestic
animals. The female fly lays her eggs in dust, sand, earth, etc., where the host is
accustomed to lie. The first stage larva, which is extremely small but very active,
on finding itself in the neighbourhood of the skin of a host, attaches itself to it, and
then bores its way into the epidermis. It now rapidly passes through its moults,
and the third stage larva, which is very distinct from the first instar in external
structure, as is the case in many of the allied CALLIPHORINAE and the OESTRIDAE,
forms a palpable tumour below the skin, its posterior extremity being in: the
vicinity of the opening into the tumour, so that air can be taken in through the
posterior stigmata. These tumours never suppurate in the ordinary course of events,
but may do so, if for any reason the larva dies.

Cordylobia (Stasisia) rodhaini, Ged.—This Calliphorine is the only other species
whose larvae are known to cause cutaneous myiasis in Africa. It appears to be
restricted to the damp forest regions of the Congo, and its larvae are normally found
under the skin of the duikers, such as the bay duiker, Cephalophus dorsalis, and
the common duiker, Cephalophus grimmt, as well in the large rodent, Cricetomys
gambianus ; man is only accidentally infested. Thin-skinned animals are the only
ones attacked, and the immobility of the host at the time of the hatching of the
larva from the egg is a necessity. As in the case of Cordylohia anthropophaga, the
eges are laid in damp earth, particularly where the hosts are accustomed to lie, and
where urine has been voided. The first-stage larva soon makes its way into the
epidermis and rapidly passes through its moults, reaching maturity in fifteen days.

A comparative study of the various larval instars with those of other
CALLIPHORINAE would be of great interest, and I trust that those who have oppor-
tunities of obtaining them will send me any specimens.

Wohlfahrtia magnifica, Schiner.—This species is the only European specific
myiasis-producing fly, and belongs to the family SARCOPHAGIDAE, or Flesh Flies.
All the species of Woflfahrtia can be distinguished from those of Sarcophaga by the
well-defined round spots on the abdomen, instead of the shimmering chequered
marks, so characteristic of all the species of Sarcophaga. The species of the allied
genus Sarcophila also have round spots on the abdomen, but the arista is only
plumose towards the base, while in the species of Wohlfahrtia the arista is plumed for
some distance. Wohlfahrtia magnifica deposits its larvae in cuts, sores, wounds,
and at the entrance to the nostrils and vagina, and on the eyelids, the fly being
attracted by an offensive discharge from such tissues or organs. It is widely dis-
tributed in Southern Russia, Asia Minor and Egypt, where it is a serious pest to
man and animals. Portchinsky has made an exhaustive study of its life-history
and habits, as well as those of some allied species. None of the other species of
Wohlfahrtia appear to be specific myiasis-producing flies.

Flies of the Family Oestridae.

In this family are included a large number of highly specialised flies, whose
larvae can only reach maturity in certain tissues and organs of special animals
and their near allies. For example, the larvae of the horse bots (Gasterophilus)
and those of the elephant bots (Cobboldia) can only live in the stomachs and duodenum
of the Equipar and the ELEPHANTIDAE, attaching themselves deeply by their man-
dibular hooks to the mucous membrane of the alimentary tract. The exact nature

244 MAJOR W. S. PATTON.

of the food of such larvae is not clearly understood, but from a large number of
dissections and examinations of the mid-guts of the larvae of the sheep nose-bot
(Oestrus ovis), I was unable to confirm Brauer’s suggestion that these Oestrid larvae
are blood-suckers. The contents of the mid-guts of larvae recently removed from
their hosts clearly suggested that they had fed on serous exudations, most probably
resulting from the irritation produced by the minute, but deep, punctures made
by their mandibular hooks in the mucous membrane. I had several opportunities
of examining the naso-pharynx of dromedaries heavily infested with the larvae of
their nose-bot, Cephalopsis titillator, and there was no doubt whatever that the mucous
membrane was in a pathological condition, and in life would exude much mucus.

The entire larval stage of the OEsTRIDAE is a long one, and is chiefly passed in
the third stage, but the adults, on the other hand, are short-lived. Sufficient food
is taken in by the third stage larva, and stored up in the form of fat-body, so that |
when the adults hatch out, particularly the female, there is ample nourishment
for the development of the eggs, and even for the hatching of the first stage larvae
in utero. The adults therefore do not feed, and as a result their mouth-parts have
atrophied for want of use. There can be very little doubt that when such larvae
are in large numbers, the host suffers, and may, as I will point out further on, even
die as a result of a massed infestation.

The OxsTRIDAE are at present grouped in a number of subfamilies according to
the structure of the third stage larva, and that of the adults. A few notes on the
species belonging to the various subfamilies may be useful to those who have
opportunities of collecting any of these flies, either as larvae or as adults. For
much of the information given below I am indebted to the exhaustive works of Rodhain
and Bequaert, as well as to those of Gedoelst and Roubaud.

1. Subfamily GASTEROPHILINAE.

All the species of this homogeneous group are myiasis-producers in their larval
stages, the larvae living in the stomach, and in one case in the duodenum, of the
horse and its allies. These larvae have occasionally been found in the stomachs of
certain carnivores which had fed on dead horses, donkeys, etc., but this infestation
is only accidental. In Russia some of the species now and then deposit their eggs on
the hair of the human face, such as the eyebrows, and the first stage larvae then pene-
trate the skin, and cause the so-called “ creeping myiasis,” the little larva burrowing
along under the skin, evidently endeavouring to find its way into the alimentary
tract; they never, however, reach the stomach, and thus do not become mature.
As would be expected, such larvae only belong to those species of Gasterophilus whose
larvae normally penetrate the skin of the horse, after hatching out of the egg, and do
not depend on moisture and friction tohelp them to leave the egg, and enter the mouth.
Gasterophilus veterinus (nasalis) and G. haemorrhoidalis are the two species whose
larvae usually cause this form of myiasis. Gasterophilus intestinalis, on the other
hand, cannot do so, as its larvae can only hatch out of the egg with the aid of friction
and moisture.

The following species of Gasterophilus are known in their larval and adult stages :—

Gasterophilus intestinalis, De Geer (equi, Clark). This is the common horse bot of
Europe and North America; it has also been recorded from many parts of Africa,
such as South Africa, the Belgian Congo, Senegal, French Guinea, the Gold Coast,
etc. It is also found in Australia and New Zealand. In India and Mesopotamia
there is a variety, bengalensis, Macq., which is very common in the north of India.
Another variety, asininus, Br., is recorded from the Anglo-Egyptian Sudan, its larvae
being parasitic in the stomach of the donkey.

Gasterophilus haemorrhoidalis, L.-—-This is also a European and North American
species, and Gedoelst records it from the Belgian Congo. Its larvae are parasitic
in the stomach of the horse.

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 245

Gasterophilus veterinus, Clark (nasalis, L.)—This species is found throughout
Europe, North America, Australia, French Guinea and Mesopotamia. Its eggs are
always attached to the hairs between the rami of the lower jaw, the fly darting
between the legs and striking the space, each time depositing an egg. Townsend
states that the eggs actually pierce the skin, but it is only necessary to examine the
ovipositor to see that this is an impossibility.

Gasterophilus pecorum, ¥.—This is the common horse bot of Russia, Hungary
and Italy ; it has also been recorded from South and West Africa. A variety, zebrae,
Rodh. & Beq., is parasitic in its larval stage in the stomach of the zebra, Equus bohm,
in East Africa.

The following species are only known in their adult stages :-—

Gasterophilus flavipes, Oliv.—This species is recorded from the Mediterranean
region, Spain, North Africa, Asia Minor and the Anglo-Egyptian Sudan. Its larvae
are believed to be parasitic in the stomach of the ass.

Gasterophilus nigricollis, Lw.—This species is only known from Bessarabia.
Gasterophilus lativentris, Brauer.——From Courland.

Gasterophilus magnicornts, Bezzi.—-This species is found in the Italian Somaliland
and is probably identical with G. intestinalis var. bengalensis, judging from the des-
cription. Its larvae are believed to be parasitic in the stomach of the horse and ass.

The following species are only known as larvae :—

Gasterophilus gedoelsti, Rodh. & Beq.—From East Africa ; the larvae are parasitic
in the stomach of the zebra, EF. béhmt.

Gasterophilus ternicinctus, Ged.—From Nyasaland; the larvae are parasitic in
the stomach of the zebra, FE. burchelli craieshayt.

There are several other doubtful larvae of Gasterophilus that have been described.
There can, I think, be very little doubt that when a critical examination is made of
fresh material of the larvae and adults of the species of this subfamily, this long list
of species will be reduced.

Rodhain and Bequaert place in this subfamily the bot-flies of the rhinoceros,
Gyrostigma, of which there are three, possibly four, good species, the adults of only
two being known.

Gvyrostigma pavesii, Corti.—This species, the adults of which are known, is parasitic
in its larval stage in the stomach of the African black rhinoceros, R. brcornis, which
is found from Abyssinia and Somaliland, through East and Central Africa, in suitable
localities down to the Cape. It is also found in the stomach of the white or Burchell’s
rhinoceros, R. simus cottont, which is found in South and South-East Africa, as far
north as the Zambesi, and again in equatorial Africa at Lado, just north of the
Equator.

Gyrostigma meruenis, Sjéstedt.—The larvae of this species are parasitic in the
stomach of the black rhinoceros, R. bicornis.

Gyrostigma conjugens, E-nderlein.—The larvae of this species are also parasitic
in the stomach of the same rhinoceros.

Gyrostigma sumatrensis, Brauer.—The larvae of this species were recovered
from the stomach of the Sumatran rhinoceros, FR. swmatrensis, which is found in
Assam and Siam, the Malay Peninsula, Sumatra and Borneo, and also from the
stomach of the Assam local race, R. sumatrensis lasiotis.

According to Rodhain and Bequaert the eggs of G. pavesii are laid on the skin
of the ears, neck and shoulders of the host, so that it is very probable that the larvae
are only able to leave the eggs when, as in the case of Gasterophilus intestinalis, they
are licked ; on reaching the mouth they can then pass down into the stomach,

246 MAJOR W. S. PATTON

2. Subfamily OESTRINAE.

This subfamily has recently been monographed by Rodhain and Bequaert, who
recognise five genera. The larvae of Oestrus, Gedoelstia and Kuirkioestrus are para-
sitic in the frontal and neighbouring sinuses of ruminants, those of RAinoestrus in
the same sinuses of equines, pigs and the hippopotamus ; and the larvae of Cephalop-
sis (Cephalomyia) in the naso-pharynx of the dromedary and camel.

The following key is taken from Rodhain and Bequaert, and may be found
useful in distinguishing the genera :—

1 (a) Wings with a section of the apical transverse nerve (the outer
section of the 4th nerve ?) turned up almost vertically to the long
diameter of the wing. Long veins 2, 3 and 4 almost of the same
length, ending before the last quarter of the wing. Transverse
marginal vein ending before the middle of the first posterior marginal
cell. Antennal pits separated on the face by a large flattened
border sig ba ee oe ae of dhe Cephalopsis.

1 (b) Transverse apical vein not vertical to the long diameter of the wing.
Transverse marginal vein ends in the middle of first posterior cell or
thereabouts a ae ae sie Be #7 se eS

2 (a) Rudiments of proboscis elongated, prolonged in front towards the end
between the palps, the extremity divided into three long cushions
which represent the labella and the rudimentary lips. Antennal pits
more or less confluent, but separated by aflattenedroundridge. Body
thick-set, covered with large papules. Abdomen short, cylindrical,
almost without pilosity, Legs short and slender. Apical transverse
nerve without an appendix - He am be Rhinoestrus.

2 (b) Rudiments of proboscis conical, not extending beyond the palps at the
base, and without traces of labellae. Venter and posterior extremity
of abdomen covered with long and fine pilosity se a £40 leds

3 (a) Antennal pits confluent, not separated by a median flattened edge.
Abdomen elongated, slightly flattened dorso-ventrally, contracted
and bent towards the base in front. Body without papules and gross
sculpture, but with short abundant pilosity, especially on abdomen ;
relatively long and robust. Apical transverse vein with an
appendix .. A oa os aus ag i Kirkioestrus.

3 (b) Abdomen squat, cylindrical, not flattened dorso-ventrally, not recurved
towards the front. Body grossly sculptured, either with tufts or
papules, less on thorax. Pilosity not very abundant. Short and
slender legs. Apical transverse vein without an appendix Sagi

4 (a) Antennal pits not confluent, separated on face by a large flattened edge.
Abdomen with denticulated tufts on dorsal surface of segments 2, 3
and 4 - - sa - = ae sts .. Gedoelstia.

4 (b) Antennal pits more or less confluent, not separated by a large flattened
ridge. Abdomen without any tufts on the dorsal surface. .. — Oestrus.

The following notes on the species give the hosts in which the larvae are found :—

Oestrus ovis, L.—The common sheep and goat bot, found wherever the hosts
occur. Its larvae are parasitic in the nasal passages, and are often present in large
numbers. Although the larvae are easily obtained, the adults are seldom seen.
The first stage larva has been recorded from man in Russia, Italy, and Algeria, in
the conjunctiva, mouth and nasal cavities ; but it is more than probable that in the
majority of these cases they were not those of O. ovis but of Rhinoestrus purpureus.

Oestrus variolus, Lw.—This is the common bot of the western hartebeest, Bubalis
major, and Jackson’s race of the Lelwel hartebeest ; its larvae are found in the nasal
passages of a large percentage of these antelopes,

~<a

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 247

Oestrus aureo-argentatus, Rodh. & Beq.—The larvae of this species are common
in the nasal passages of the roan antelope, Hippotragus equinus, and its local races,
and in the western and Jackson’s hartebeest.

Oestrus macdonaldt, Ged.—The larvae of this somewhat rare species have so far
only been found in the nasal passages of Lichtenstein’s hartebeest, Bubalis
lichtensteint.

Oestrus interruptus, Ged.—The mature larva of this species is parasitic in the
nasal passages of Coke’s hartebeest, Bubalis cokei, the brindled gnu, or blue wilde-
beest, Connochoetes taurinus johnstont, and the bastard hartebeest, Damaliscus
lunatus.

Oestrus disjunctus, Ged.—The larva of this species is parasitic in the nasal
passages of the bastard hartebeest, and the roan antelope.

Oestrus compositus, Ged.—The larva of this species is found in the nasal passages
of Lichtenstein’s hartebeest.

Rhinoestrus purpureus, Brauer.—This species, the larvae of which are parasitic
in the nasal passages of the horse, the mule and zebra in Europe, Asia Minor, Palestine
and Africa, is an important Oestrid, as the female fly has the habit of depositing
its larvae in, or near, the human eye, and may thus lead to the loss of an eye. It is
probably the larvae of this species that has been confused with that of Cestrus ovis
in those cases in which the latter has been recorded from the human face.

Rhinoestrus phacochoert, Rodh. & Beq.—The larva of this rare species is found
in the nasal sinuses of the warthog, Phacochoerus aethiopicus, in the Belgian Congo ;
the adult fly is not known.

Rhinoestrus mivarletit, Rodh. & Beq.—The larvae of this species are parasitic
in the nasal sinuses of the red river-hog, Potomochoerus porcus, in the Belgian Congo
and adjacent parts of West Africa; the adult flv is not known.

Rhinoestrus hippopotami, Griinberg.—The larvae of this interesting Oestrid.
the only species known from the hippopotamus, is parasitic in its nasal cavities.

Gedoelstia cristata, Rodh. & Beq.—The larvae of this species are found in the
nasal cavities of various antelopes such as Lichtenstein’s hartebeest, the western
hartebeest, Jackson’s variety of the Lelwel hartebeest, the white-bearded race of
the blue wildebeest, and the topi or variety of the Senegal hartebeest.

Gedoelstia hassleri, Ged.—The larvae of this species are parasitic in the nose and
accessory sinuses of Lichtenstein’s hartebeest and the bastard hartebeest.

Kirkioestrus surcoufi, Ged.—-The larvae are parasitic in the nasal cavities of the
western hartebeest, Jackson’s hartebeest, and species of the lechwe in various parts
of West Africa.

Kirkioestrus blanchardi, Ged.—Its larvae are parasitic in the nasal cavities of
Lichtenstein’s hartebeest in the basin of the Zambesi.

Kirkioestrus minutus, Rodh. & Beq.—The larvae of this species are parasitic in
the nasal cavities of Jackson’s hartebeest and the topi, in the Congo.

Cephalopsts titillator, Clark (Cephalomyia maculata, Macq.).—The larvae of this
species, the only bot-fly of the CAMELIDAE, is parasitic in the naso-pharynx of the
_ dromedary and the camel, wherever these animals are found.

3. Subfamily COBBOLDIINAE.

The stomach-bots of the Indian and African elephants are placed in this
subfamily, and there seems little doubt that they are near allies of the equine
stomach-bots, the GASTEROPHILINAE.

248 MAJOR W. S. PATTON.

Cobboldia elephantis, Steel—The larvae of this species are parasitic in the stomach
of the Indian elephant. I have been told by those who have to do with the elephants
kept by the Government of Madras, in various parts of South India, that this bot-
fly is a serious pest. There seems little doubt that when the larvae are in large
numbers in the stomachs of the elephants, especially the young animals, they suffer
in health, often being on the sick list with digestive troubles, and are unable to
work; and in bad infestations the young animals may even die. Mr. Chari,
Veterinary Inspector, Mount Stuart, sent me a large number of third stage larvae
of C. elephantis that were collected from the stomach of a calf elephant, which was
so full of larvae that there was hardly any room for a pin to pass into it. The eggs
are laid at the root of the tusk or trunk, and the larvae only hatch out when the
eges are moistened. I had a large number of these eggs collected about a month
after they were laid, and as soon as they were placed in some water the young larvae
immediately burst open the eggs and emerged ; they are very active and can crawl
considerable distances.

The African elephant is parasitised by the larvae of three species of Cobboldia.
The larvae of Cobboldia loxodontis, Brauer, have been recorded from Uganda, the
Belgian Congo, and the Ivory Coast, but its distribution is probably much wider ;
those of C. chrysidiformis, Rodh. & Beq., have so far only been found in the stomachs
of elephants from the Belgian Congo; while those of C. parumspinosa, Ged., have
been recorded only from the basin of the Zambesi.

4. Subfamily HyPODERMATINAE.

This subfamily contains all those OESTRIDAE whose larvae cause cutaneous
myiasis in ruminants, chiefly cattle and antelopes. The following are the more
important species :—

Hypoderma bovis, De Geer.—The larvae cause the well known warbles, which
are common in the hides of cattle in Europe and North America. Captain Cross,
C.V.D., has sent me what appear to be the larvae of this species from the skins of
goats in the Punjab, a very large percentage of the animals being infested.

Hypoderma lineata, de Villers.—The larvae are also parasitic in the hides of
cattle in Europe and North America, but are not so common as those of H. bovis.
It is only recently that the method of infestation has been worked out in the case
of Hypoderma bovis by Carpenter in Ireland, Glaser in Germany, and Hadwen in
Canada. The females of these two species lay their eggs on the hairs of the legs
of cattle, and the larvae on hatching out penetrate the skin adjacent to the hair,
and then slowly migrate upwards until they reach the submucous coat of the
oesophagus, resting there for some time in the second larval stage. Later they
pass along the muscles of the back and eventually reach the back and sides of the
animal, where they make a small opening in the skin. When mature they leave the
skin and pupate in the ground.

Hypoderma diana, Brauer, and H. actaeon, Brauer.—The larvae of these two
species are parasitic in the skins of the red deer in Germany and Britain.

Hypoderma silenus, Brauer, is parasitic in its larval stage in the skin of the ass
in Egypt: Hypoderma desertorum, Brauer, is also found in Egypt, but its host is
not known. Hypoderma gazellae, Ged., is parasitic in its larval stage in the skin
of Grant’s gazelle in East Africa. Hypoderma corinnae, Crivelli, in the skin of
the Dorcas gazelle, Gazella dorcas.

Oedemagena.—This. genus, which contains only one species, differs from
Hypoderma in having small oval labella, which are absent in Hypoderma. Oede-
magena tarandi, L., is parasitic in the hides of reindeer in Lapland and Norway.

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 249

5. Subfamily CUTEREBRINAE.

In this subfamily are placed a somewhat heterogeneous collection of genera and
species, all the larvae of which are parasitic in the skin and subcutaneous tissues
of small rodents, chiefly MurtDAE ; they are only found in North and South America.
Townsend raises the subfamily to a family, the CUTEREBRIDAE, and gives the following
key to the genera :—

(1) No facial carina; antennal pit large and deep; antennae elongate.. 2.
Facial carina present ; antennal pit small and shallow ; antennae short — 3.
(2) Epistoma rather broad, projected obliquely forward and downward
between the peristomalia; arista thickly long-plumose to tip

Pseudogametes.

Epistoma very narrow, projected straight downward between the
peristomalia ; arista with hairs on upper side only .. Dermatohia.
(3) Arista nude Rogenhofera.

Arista with hairs on upper side and on apical part of lower side .. 4.
(4) Antennal pit extended below in a tapering prolongation, the peris-
tomalia converging obliquely and meeting near the lower end of
carina ff Ho ys a .. Cuterebra.

Antennal pit circumscribed below, subcircular, the peristomalia
parallel and closely approximated to the carina for about the lower
half of its length .. e 3 oe wr 7 .. Bogeria.

The following are the more important species :—
Pseudogametes hermanni, Br., and P. semiatra, Wied., from the subcutaneous
tissues of MuRIDAE in Brazil.

Dermatobia hominis, L.—The larvae of this species are chiefly found in the skin
of cattle in tropical America and the neighbouring islands; man is seldom parasitised.
It is now known that the female fly lays its eggs in small batches on the bodies of
blood-sucking and sweat-loving insects, such as the Culicid, Janthinosoma lutzt, and
species of Anthomyia, and the larvae hatch out when the egg-carrier visits the

vertebrate host to feed, penetrating the skin and forming a local tumour.

Rogenhofera grandis, Guérin, R. trigonocephala, Br., and R. dasypoda, Br., from
the skins of MurIDAE in Argentine and Brazil. Cuterebra americana, F., C. cunicult,
Clark, C. analis, Macq., C. approximata, Walk., C. hisirio, Coq., C. tenebrosa, Coq.,
C. atrox, Clark, and C. maculosa, Knab, all from the skins of small rodents in North
and Central America.

Cuterebra ephippium, Latr., larvae parasitic in the skins of MurtpaE in French
Guiana, C. patagona, Guér., from Patagonia, C. megastoma, Br., from South America,
C. funebris from Trinidad, C. apicalis, Macq., C. cayennensts, Macq., C. rufwentris,
Macq., C. nigrocincta, Aust., C. sarcophagoides, Lutz, C. nigricans, Lutz, C. tnfulata,
Lutz, and C. schmalzi, Lutz, from Brazil. Bogeria emasculator, Fitch, B. grisea,
Coq., B. buccata, F., B. fontenella, Clark, B. princeps, Aust., B. fasciata, Swenk.,
B. scudderi, Towns., all from North and Central America.

From the studies of Hadwen, and Parker and Wells, it seems that these Cutereb-
tine bot-flies deposit their eggs on the hairs of their hosts, and that they are then
licked off and enter the alimentary tract, later migrating outwards to the skin, where
they cause dermal myiasis.

6. Subfamily CEPHENOMYINAE.

In this subfamily are included a small number of Oestrids, the larvae of which
are parasitic in the nose and accessory sinuses of the CERVIDAE and BoyrpaE and one
species in the oesophagus of the African elephant. Most of the species are found in

250 MAJOR W. S. PATTON.

Europe and in North and South America. As in the case of the OESTRINAE, the
females are viviparous and deposit their larvae at the entrances of the nostrils, the
larvae then migrating up into the frontal sinuses, and down, even into the oesophagus.

Cephenomyia auribarbis, Mg., is a common parasite in its larval stage in the
red deer in Europe; the larvae of C. aricht, Br., in the elk ;. C. irompe, L., ansthe
reindeer ; C. stimulator, Clark, in the roe; and C. abdominalis, Aldrich, from the
Adirondacks, New York. C. pratti is parasitic in its larval stage in the American
elk, the mule deer, and several other deer in North America, C. phobifer is only
known in its adult stage from North America, and C. macrotis has been recorded
in its larval stage from the mule deer in North America.

Semi-specific Myiasis-producing Diptera.

In this group I include all those flies which, though normally breeding in the
bodies of dead animals, and even in vegetable matter, will occasionally lay their eggs,
or deposit their larvae, in the diseased tissues of man and animals. The female fly
is in each case attracted by a foul discharge from a sore, wound or diseased organ,
such as the nose or ear, or even in soiled wool. The following species belong to this

roup :—
eas Calliphora erythrocephala, Mg.
: vomitoria, L.
ui quadrimaculata, Swed.
, (Anastellorrhina) augur, L.
Cochliomyia (Chrysomyia) macellaria, L.

. viridula, R.D.
Chrysomyia megacephala, F. (dux, Esch.).
= marginale, Wied.
" albiceps, Wied. (rufifacies, Guérin).
isp varipes, Macq.

Pollenia stygia, F.
Phormia regina, Mg.
Cynomyta cadaverina, R.D.
Lucilia sericata, Mg.
6 caesar, L.
A. areyricephala, F. (serenissima, Macq.).
% lasmaniensis.
Sarcophaga ruficornis, F.
a chrysostoma, Wied.
7 plinthopyga, Wied.
Aphiochaeta xanthina, Speiser.
., rufipes, Mg.
A few notes on the above species may be useful.

Calliphora.—-All the species belonging to this genus of blowflies are large robust
insects of a dark blue colour, often with lighter patches on the abdomen. The palpi
are nearly always reddish, the legs black, and the thorax adorned with large bristles.
The females, which are oviparous, occasionally lay their eggs in living tissues of man
and animals, though normally ovipositing in decaying animal matter ; the habit of
ovipositing in diseased tissues is not well established in the species of this genus.

Cochliomyia.—These blowflies can be distinguished at once from their Old World
allies of the genus Chrysomyia (Pycnosoma) by the well-marked dark dorsal
thoracic stripes. C. macellaria, the notorious screw-worm fly of tropical America,
like its ally, Chrysomyia bezziana, will lay its eggs in any diseased tissues, but unlike
it, will also breed freely in the bodies of dead animals, and even in decaying vegetable
matter. Its control is therefore a much more hopeless task than that of Chrysomyia

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 251

bezziana. The larvae of Cochliomyia viridula are known to cause cutaneous myiasis
in man and animals in British Guiana and Trinidad. Doubtless there are several
other species of this genus in South America with like habits.

Chrysomyia (Compsomyia, Pycnosoma).—The blowflies of this genus are usually
of a bluish green colour, and have well-marked horizontal bands on the hind borders
of the abdominal segments. The thoracic bristles are reduced, the dorsal surface being
covered instead with fine hairs. Chrysomyia megacephala is one of the species that
occasionally deposits its eggs in the tissue of animals in India. It is very similar
in general appearance to Chrysomyia bezziana, as pointed out above. Chrysomyia
marginale is acommon African species, and though mainly breeding in decomposing
bodies of animals and birds, will occasionally lay its eggs in, or near, diseased tissues.
Chrysomyia albiceps (rufifacies) is, according to Froggatt, a notorious sheep maggot
fly in Australia, its larvae being found in company with those of two other blow-
flies, Anastellorrhina augur and Pollenia stygia. In India I have noted that the
larvae of albiceps are entirely predaceous, feeding on the larvae of other CALLIPHOR-
INAE, so that it seems possible that it lays its eggs in soiled wool in Australia, being
attracted by the presence of other larvae ; this point requires further investigation.
Chrysomyia varipes is a smaller species with a somewhat similar larva; it also
breeds in soiled sheep wool in Australia.

Anastellorrhina augur is a well-known sheep maggot fly in Australia. It is a large
species, suggesting affinities with Calliphora. The larva, as I will point out further
on, is very similar in structure to that of Calliphora erythrocephala.

Pollenia stygia (villosa) is another allied species, which is placed in this genus
because of the fine yellow hairs on the sides of the thorax. It is a serious pest to
farmers in Australia.

Phormia regina, though a common fly in Europe, is not, so far as I am aware,
known to cause myiasis in animals. In America, however, it often lays its eggs in
soiled wool and in wounds of sheep, particularly old suppurating wounds. Bishopp
states that its larvae are often found in sheep with broken horns.

Cynomyia cadaverina very occasionally causes myiasis, more especially in very
foul wounds on animals. It is, however, a regular breeder in decaying animal matter.

Lucilia.—The species of this genus of bright metallic green blowflies can be dis-
tinguished from the species of the genus Chrysomya by noting that there are many
strong bristles on the thorax, arranged in two parallel rows. Luctlia sericata is the
most notorious species, for it regularly lays its eggs in the soiled wool and tissues of
sheep in Europe, and more particularly in Scotland. It is also recorded from
Australia as a sheep-maggot fly. Luctlia caesar only very occasionally lays its eggs
in living tissues. Lucilia argyricephala is the common Oriental species and occasion-
ally lays its eggs in the diseased tissues of animals. Lucilia tasmaniensis is said
to be a myiasis-producing species in Tasmania.

Sarcophaga.—All the species of this cosmopolitan genus are very characteristic
large grey flies with red eyes. Though easily recognizable generically the species
are exceedingly difficult to determine. All are semi-specific mviasis-producing flies
normally breeding in dead bodies and decomposing animal matter. In India there
is, at least, one species which deposits its larvae in wounds, particularly those exuding
very foul pus, and the larvae, owing to their large size, cause very extensive destruc-
tion, so that there can be no mistaking a case of myiasis caused by such larvae. This
species is believed to be vificornis, but this identification is not certain. I would
particularly urge all those who have opportunities of collecting the larvae of species
of Sarcophaga from cases of myiasis to breed out the flies, for it is only by doing this
that we can be certain of the species and can then study its habits and distribution.
Sarcophaga lambens, Wied., and Sarcophaga pyophila, Neiva, are recorded as causing
myiasis in Central and South America. Sarcophaga chrvsostoma, Wied., in British
Guiana, and S. plinthopyga, Wied., are also notorious myiasis-producing species.

252 MAJOR W. S. PATTON.

A phiochaeta.—The species of this Phorid genus can be recognised by their peculiar
habit of moving in a jerky manner, and by the structure of the antennae and the charac-
teristic venation. A. xanthina (ferruginea) is a yellowish brown species, which 1s
widely distributed in the tropics, and its larvae cause cutaneous myiasis in man and
animals. A. vifipes is also a myiasis-producing species, but not so common as
xanthina.

Accidental Myiasis-producing Flies.

In this group are included an assemblage of unrélated Diptera—the larvae of which
occasionally find their way into the intestinal tract of man. They normally breed
outside the human body in organic and vegetable matter, some of which is used as
human food, and people who are not particular with regard to what they eat become
infested. It should be clearly understood, however, that these flies do not intention-
ally deposit their eggs or larvae in certain human foods in order that their larvae may
reach the intestinal tract ; the food is merely their usual breeding ground. Although
many of the OrsTRIDAE live in their larval stages in the intestines of animals, they are
not accidental myiasis-producing flies, the intestinal tract is the only place in which
their larvae can reach maturity.

So far as I can gather, intestinal myiasis is an uncommon condition, even in
countries like India, for in spite of the fact that I specially drew attention to it in
the circular mentioned above, which was sent to every medical and veterinary officer,
and in spite of the fact that some medical men have told me that it is very common in
parts of the country, I received such larvae from only two cases.

The well known “ rat-tailed”” larvae of Evistalis tenax and Helophilus pendulus,
which normally live in foul ditch water, may be acquired by those, particularly
children, who drink such water, when they are likely to swallow large numbers of
small larvae of these drone-flies. In the same way, vegetables, such as lettuces and
mustard and cress, grown under insanitary conditions and eaten without being washed,
may contain eggs and larvae of Ervistalis tenax and allied species. The eggs and
larvae of Fannia and Anthomyia may also be swallowed intact in uncooked vegetables,
especially raw carrots. Onion fly larvae may be swallowed when eating diseased
spring onions, and species of Drosophila in over-ripe fruit, such as bananas and oranges.
Sarcophaga larvae are sometimes swallowed in imperfectly cooked meat, particularly
cold meat, and game that is commencing to decompose. In the case of Aphtochaeta
xanthina (ferruginea), the larvae of this small Phorid are most probably swallowed in
meats of all kinds, especially when beginning to decompose, as this substance is a
favourite breeding ground for the fly. One remarkable case has been published in
which the larvae, puparia and adults of this fly were passed in the faeces of a patient
for a considerable time, suggesting that the larvae were able to reach maturity in
the intestine, and the puparia remain alive, so that the adults could hatch out.
Recently in Coonoor, I attempted to infect monkeys by feeding them on large numbers
of eggs and larvae of this species, but although a most careful search was made for any
stages in their excreta, I was never able to recover either larvae, puparia or adults.
It would be worth while repeating this experiment, using only one monkey and making
a more thorough search than I was able to undertake. Such small larvae as those of
Aphiochaeta easily escape detection when the tray under such a monkey contains a
considerable quantity of remains of food in addition to its faeces.

So far as I am aware, the larvae of the CALLIPHORINAE are unable to live in the
intestine of man and animals, but in order to set aside any doubt on the subject I fed
a dog with large numbers of eggs and larvae of some common Indian species, but
not a single living larva was passed out, and it would appear that such larvae, when
swallowed, soon die. But during the course of my enquiry into the myiasis-
producing flies of India, two masses of eggs of Chrysomyia megacephala were sent
me as having been passed out in the faeces of a patient. The eggs were intact, and

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 253

the medical officer was emphatic in his statement that they were actually passed out,
and had not just been deposited on the faeces. In any case, Chrysomyia megacephala
certainly does not breed in human excrement, so far as I know. It would be worth
while carrying out some more experiments in this direction to see whether the eggs of
CALLIPHORINAE can hatch out in the intestine, and the larvae reach maturity and be
passed out in a living condition.

Lastly, it is necessary to refer to the larvae of Fannia canicularis from the urinary
tract. There is at present no satisfactory explanation as to how such larvae can reach
the human bladder. The usual explanation that they pass down the urethra when
very small, seems to me to be highly improbable, at least in the case of the male
urethra, for it will be remembered that the larva of Fannta, even when small, is not
entirely smooth, and would cause very considerable irritation. The presence of such
larvae in this organ is at present a mystery, and the subject requires to be
re-investigated.

Identification of the Larvae of Myiasis-producing Diptera.

It will be remembered that the larvae of the higher Diptera are soft cylindrical
maggots, usually of a grevish white to yellowish white colour. The head end is
pointed and the body gradually increases in diameter backwards, the posterior end
being truncated, with a concave surface looking upwards and backwards. Such a
larva is divided into segments, the number of which is at present a matter of dispute,
and it is not possible to homologise them with those of the adult. The head contains
a characteristic skeleton consisting of a number of sclerites, spoken of as the cephalo-
pharynx. The mouth is armed with a pair of mandibular hooks, and the anterior
spiracles open at the sides of the apparent third segment. The posterior stigmata are
placed on the concave surface of the apparent eighth abdominal segment. Banks,
MacGregor and Metcalf have drawn attention to the value of the posterior stigmata
in determining the larvae of the higher Diptera. Metcalf in particular says that,
“So superior are the characteristics drawn from this part of the larva, that I regard
most descriptions which omit consideration of them as practically worthless. For
in most cases the general features of shape, colour, etc., may be found to apply almost
equally well to other species. The most available, absolutely diagnostic characters
are to be found on the posterior stigmata.’’ In my own studies of the larvae of Diptera
I have found that the characters of the posterior stigmata, cephalopharynx and
anterior spiracles, taken together, afford the best and only reliable means of accurately
determining the larva of any species of Diptera. and more particularly of those
of the higher Diptera. In making a preparation of the posterior stigmata of a Muscid
larva for purposes of identification, I always make it a rule to include the cephalo-
pharynx and anterior spiracles as well. 1 will now shortly describe these structures
in the various types of larvae of the myiasis-producing Diptera, and point out how
their characters can be utilised in determining any particular larva.

Postervor stigmata. In the third-stage larvae of all the higher Diptera the posterior
stigmata consist of a pair of chitinous plates, which, when cleared in caustic potash,
are seen to be surrounded by a ring of chitin, usually spoken of as the peritreme.
Situated within this ring there are three or more openings which lead into the tracheae.
Each opening is guarded by a number of delicate chitinous rods, giving the
appearance of a grating; these rods form a network across the inner side of the
openings. Plate 5, fig. 1, represents a vertical section through one of the stigmatic
plates of the third-stage larva of Pollenia stygia. The deeply shaded portions
represent the chitinous parts of the plate, and the lighter-shaded parts the epidermis.
The three slits open into a large cavity, to the margins of which the trachea is attached ;
Lowne termed this cavity the vestibule. Just inside each slit there is a series of
branched yellow chitinous rods, known as the grating, and lining the whole cavity
a delicate network of epidermal fibres. The grating is evidently of use in preventing

(4183) =

254 MAJOR W. S. PATTON.

the entry of foreign particles into the vestibule, and the epidermal fibrous network
probably aids in retaining the air in the vestibule. The tracheae are lined externally
by characteristic peritracheal cells. Situated on the inner and lower side of each plate,
and usually within the chitinous ring, there is a small raised area, spoken of by Banks
as the “ button.” In a vertical section through this structure it will be noted that it
is a fine channel into the vestibule and is lined by a layer of peritracheal cells; so far as
I can ascertain, it is an accessory opening, but I am unable to say at present whether
it is functional or not. It is certainly not a point of attachment for muscles, as it
is within the vestibule. This short description will enable the reader to understand
this wonderful bit of apparatus. All the types which I have sectioned are of similar
structure, no matter whether the tracheal slits be straight, as in the CALLIPHORINAE,
curled as in Musca, or multiple as in Oestrus and allied genera, so that I can see little
use in splitting them up into various types according to the shape of the slits. The
important point to be ascertained is whether the straight or the curled shit is the more
primitive, and this can only be ascertained by a comparative study of the stigmatic
openings of a large number of the first stage larvae of the higher Diptera. An oval
opening, such as is present on the anterior spiracles of most Dipterous larvae, seems
to be the most primitive type, and from such an opening the various slits seen to-day
in the posterior stigmata of the higher Diptera may have originated, but this is at
present mere conjecture.

Turning now to the various kinds of slits, leading into the tracheal vestibule of
the posterior stigmatic openings, we note the following types :—(1) Curled slits,
(2) straight slits, and (3) round or oval openings.

1. Cuwrled slits. This type of opening is characteristic of the larva of Musca and
allied genera, the three openings occupying an excentric position on the plate, and
the ends of the upper and lower appearing to coalesce with those of the middle slit.
The button area is well within the chitinous ring, which, in this case, is often
D-shaped and broad. In Platev, figs. 2,3, 4 and 5, are illustrated the posterior stigmata
of Musca domestica, Musca nebulo, Musca humilis and Philaematomyia crassirostris,
and without going into details it will be seen that, though very closely similar
to each other, it is possible to note small distinctions sufficient for determining the
species. But I would like to point out that the determination is considerably
facilitated by comparing the structure of the cephalopharynx and anterior spiracles
of each species at the same time. Recently I was able to determine the larva of
Musca (Philaematomyia) crassirosivis sent me from a case of intestinal myiasis by
comparing the posterior stigmata with those of Indian species of Musca. Curled
slits are also seen in the posterior stigmata of the larvae of the GASTEROPHILINAE,
especially in the larvae of the species of Gyrostigma. Here, however, the grating is
simpler than in the larvae of Musca.

2. Straight slits. This type of slit is characteristic of the opening of the posterior
stigmata of the larvae of the CALLIPHORINAE, SARCOPHAGIDAE and some OESTRIDAE,
such as the larvae of Cobboldia. In the larvae of the CALLIPHORINAE the posterior
stigmata are of two kinds, one typically seen in the larvae of Calliphora and Lucilia,
in which there is no break in the chitinous ring and the button is enclosed by it ; the
other in Chrysomyia and Cochliomyia, in which the ring of chitin is so thin on the inner
and lower angle of the plate as to give the appearance of a’break, the button lying in
what is but a thin layer of chitin. This type is illustrated in Plate vi, figs. 9, 10
and 11, which represent the posterior stigmata of Calliphora erythrocephala, Lucilia
sericata and Lucilia argyricephala, all drawn to the same scale. Further, it should
be noted that in each of these there are characteristic breaks in the chitin, between
and close to the slits, appearing as round or oval spaces ; these clear spaces are
extremely constant in the same larva. This close similarity between the posterior
stigmata of Calliphora and Lucilia makes it difficult to separate the larvae of Calli-
phora erythrocephala from those of Lucila sericata. Veterinary officers may find

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 250

this so, when they have to determine a larva taken from a sheep in this country.
A careful examination of the plates when cleared in caustic potash and mounted on
a slide will, however, help to separate the species. In the larva of Calliphora erythro-
cephala, as well asin that of C. vomitoria, the plates are distinctly larger and further
apart than those of the larva of Lucilia sericata. In addition, in Calliphora there are
two processes of the rim which project in between the upper and middle slit, and the
middle and lower slit, whereas in Lucilia sericata there is only one projection between
the middle and lower slit. These points, together with certain differences between
the cephalopharyngeal skeletons and anterior spiracles of the two species, will make
identification easy, but it is necessary to point out that mere examination of the
larvae with a hand lens is not sufficient. Plate vi, fig. 13, illustrates the posterior
stigmata of the third-stage larva of Anastellorrhina augur, and it will be noted that
they are very similar to those of Calliphora. This alone, in my opinion, indicates
the affinity of this blow-fly. The larva of Cynomyta cadaverina also has posterior
stigmata of this type, as is seen in fig. 9; here, however, the clear spaces are not seen.
The posterior stigmata of Lucilia argvricephala are smaller than those of L. sericata
and more widely separated.

In the larvae of Chrysomyia and Cochliomyia the posterior stigmata appear to
have a break in the chitinous ring on the lower and inner face, the plates are, as a
whole, larger, and the button area is situated at the lower angle. Plate vi, figs. 14,
15, 16 and 17, illustrate the posterior stigmata of the larvae of Chrysomyia mega-
cephala, Chrysomyia bezziana, Cochliomyra macellaria and Phormia regina, which also
belongs to this group. It is interesting to note that in the case of the two myiasis-
producing species, macellaria and bezziana, the slits are short and wide, those of
bezziana having characteristic clear spaces between and near the slits; these two
flies are evidently very closely related.

Apart from other differences, such as the shape of the truncated posterior end,
the larvae of the SARCOPHAGIDAE can always be distinguished from those of any of
the CALLIPHORINAE by noting that the posterior stigmata are situated in a deep cleft,
and that the internal slit slopes downwards and outwards, the middle almost
straight down, and the outer slit a little inwards and backwards. The
plates are always large and the slits long and somewhat narrow, and there is a
characteristic break in the chitinous ring at the inner and lower angle ; the posterior
stigmata of the third stage larvae of a species of Sarcophaga taken from a case of
cutaneous myiasis in India are shown in Plate vi, fig. 18. The cephalopharynx and
anterior spiracles of the larva of Sarcophaga are also very distinct, as I will point out
further on, so that there should never be any difficulty in identifying a larva of this
family of flesh-flies.

The posterior stigmata of the larvae of the elephant bots are very characteristic.
The plates have here coalesced and the chitinous ring is feebly developed, the margins
of the ring being probably represented by an island of chitin on the inner face.
The slits are long and almost straight, situated close to one another; fig. 6, Plate v,
illustrates the posterior spiracles of the third-stage larva of Cobboldia elephantis from
the Indian elephant. The button area has apparently disappeared in this type of
posterior stigmata. On larval structure alone, then, Cobboldia suggests affinities with
a Calliphorine on the one hand and a Sarcophaga on the other.

3. Single vound or oval openings. This type of posterior stigmata is well seen in
the third-stage larva of the sheep nose-bot, Oestrus ovis. The plates are large, some-
what kidney-shaped, and the whole surface is dotted with minute openings of varying
shapes and sizes. The button area is well marked and almost central in position.
A similar plate is seen on the larva of the camel bot, Cephalopsis titillator, Plate v,
fig. 7; the button area here is situated about the middle of the inner face of the plate.
and is somewhat difficult to see. The posterior stigmata of the larva of Hypoderma
(Plate v, fig. 8) also belong to this type, the openings here being arranged in more

(4183) 1) 2

256 MAJOR W. S. PATTON.

or less regular rows, with strong bars of chitin projecting in between them. This type
of posterior stigmata would, at first sight, appear to be extremely complicated, but I
am inclined to think that they merely represent a breaking up of the original long slit
into smaller ones to allow of a larger amount of air being taken in. The origin of
this type can, however, only be determined by a comparative study of the first-stage
larvae of a number of different species.

Cephalopharynx. This structure in the third-stage larva of the higher Diptera

consists of a number of paired sclerites, some of which have fused, articulating with
each other to form a strong chitinous skeleton. At the anterior end there is a pair of

Fig. 1.—Cephalopharyngeal skeletons of third-stage larvae
of: (a) Musca nebulo ; (b) Calliphora erythrocephala ;
(c) Lucila sericata; (d) Sarcophaga sp.
mandibular sclerites or hooks, each with a broad base, which project between the oral
lobes and are used by the larva to attach itself to any object, and also in progression.
Articulating with the ventral surface of the base of each hook there is a small,
irregular sclerite, which Hewitt speaks of as the dental sclerite. Posteriorly the base
of each mandible articulates with an irregularly shaped sclerite, which consists of two
parts fused ventrally. These, again, articulate with small processes on the anterior
end of the large pharyngeal sclerite, which at once suggests the fulcrum of the adult
fly, and is shaped like the old-fashioned Spanish stirrup-iron.

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 257

The shape of the mandibular sclerites and the fulcrum (pharyngeal sclerite) is
very characteristic in the third stage larvae of the different genera of the higher
Diptera. In fig. 1 (ad) the cephalopharyngeal skeletons of the third-stage
larvae of Musca nebulo, Calliphora erythrocephala, Lucilia sericata and Sarcophaga
are illustrated, all being drawn to the same scale. The differences are obvious,
and therefore I do not intend here giving a detailed description of them. It is, how-
ever, necessary to point out that in the case of the larvae of C. erythrocephala and
Lucilia sericata the differences are small, but nevertheless sufficient to determine the
species. I will on another occasion deal at length with the structure of the cephalo-
pharyngeal skeletons of a number of the larvae of the higher Diptera. I would,
however, like to point out that in the larva of Musca and allied genera there are always
two mandibular hooks, and not one, as is usually stated in the books and papers on the
subject. This error is no doubt due to the fact that the left hook is shorter and
narrower than the right, and is therefore easily overlooked. It is a valuable diagnostic
character in this group.

Anterior spiracles. The anterior spiracular openings are always situated at the
sides of the lower border of the apparent third segment. Each consists of a fan-shaped

Fig. 2.—Anterior spiracles of third-stage larvae of: (a) Musca
domestica; (b) M. nebulo ; (c) Calliphora erythrocephala ;
(d) Lucilia sericata; (e) Savcophaga sp.

chitinous structure with a varying number of small finger-like processes at the apices
of which there are oval openings. These openings lead into a miniature vestibule,
which is continuous with a large trachea ; in the majority of the third-stage larvae the
anterior spiracles are functional. They are always absent in the first-stage larva,
and appear only in the second instar. Although there is great variation in the number
of the finger-like projections in the same species, and even in the same larva, these
structures are very useful for diagnostic purposes when taken along with the charac-
ters of the posterior stigmata and the cephalopharyngeal skeleton. In the third-
stage larva of Musca domestica (fig. 2, a) there may be five, six, or seven processes ;
in M. nebulo (fig. 2, b) seven or eight; in Calliphora erythrocephala (fig. 2, c) eleven or

258 MAJOR W. S. PATTON.

twelve: in Lucilia sericata (fig. 2, d) seven or eight. In Sarcophaga (fig. 2, e) the anterior
spiracle is large, and there may be sixteen or more processes, and the vestibule has
many small, clear circular areas.

I have, I think, now shown that if the characters of the posterior stigmata,
cephalopharyngeal skeleton and anterior spiracles are taken together, the species to
which any given larva belongs can be determined with certainty.

The following key to the larvae of some of the more important myiasis-producing
Diptera (exclusive of the OFSTRIDAE) will be useful for Telerence:

Key for Identification of some of the Larvae of the M yiasis-producing Diptera.

1. Larvae with fleshy processes ; posterior spiracles at the end of tubercles or a
tube.

(a) Small broad, dirty white larvae about 4mm. in length, with small pointed
fleshy processes in pairs on dorsum, increasing in length from before back-
wards ; two long fleshy processes at sides of last segment, which is the
broadest; posterior stigmata on brown chitinous tubercles, each with a
narrow opening; movements caterpillar-like

Aphiochaeta xanthina (ferruginea).

(b) Similar larvae, with hairy processes. as i -« ALryupipes:

(c) Medium-sized to large larvae with long pointed fleshy processes dorsally and
laterally, increasing in length from before backwards, with small spines at
their bases, shaft and ends bare; body compressed dorso-ventrally ; posterior
stigmata stalked, consisting of four lobes, each with an opening situated on
dorsum of sides of apparent 8th segment re .. Fannia canicularis,

(d) Fleshy processes with numerous branches, giving the process a feathery
appearance fe ss - - - Ay Fannia_ scalarts.

(ce) Medium-sized to large larvae with long pointed fleshy processes with a small
tuft of spines at their apices ; posterior stigmata in cleft at end of apparent
8th abdominal segment, consisting of large chitinous plates with broad rims,
and three straight slits directed downwards and inwards

Chrysomyia albiceps (rufifactes), C. varipes.

(f) Large soft larvae ; integument covered with hairs and spines ; seven pairs

of short pseudopods on ventral surface ; posterior stigmata situated at the

end of a long tube, which can be telescoped or extended at will
Eristalis, Helophilus.

1/. Medium-sized to large smooth larvae without any fleshy processes ; posterior
stigmata normal, not at the end of tubercles or a tube :

2. Posterior stigmata somewhat D-shaped ; button area situated well within
the ring; slits curled.

(a) All slits with three convex loops outwards .. Musca domestica and allies.
(b) Upper and lower slits with two loops, convex externally ; middle slit with
OnbyOne gis ays ahs ab as the .. Phulaematomyia.

2/. Posterior stigmata large, rounded or oval, with straight slits bs rege

3. Chitinousring complete andinclosing button area. Calliphora, Luctlia, Cynomyta.

(For distinctions see drawings of posterior stigmata.)

3/. Chitinous ring incomplete at AAs 7. he et te cde oA

4. Slits all directed downwards and inwards. Chrysomyia, Cochliomyia, Phorma.
(For distinctions see drawings of posterior stigmata.)

4!, Internal slits comma-shaped, directed downwards and outwards ; middle

slits almost straight down ; outer slits directed downwards and inwards
Sarcophaga.

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 259

I will now suppose a medical or veterinary officer has a case of myiasis, and wishes
to know how best to set about rearing the larvae, breeding out the flies and identifying
at least the genus to which the larva belongs. If the case is one of dermal or subdermal
myiasis, for instance a cut, sore or wound on the human body or that of an animal,
there should be no difficulty in obtaining a number of living larvae. It is sometimes
quite easy to extract the larvae without injuring them, but in the case of Chrysomyra
bezziana the strongly developed, backwardly directed spines catch in the tissues,
and, as often as not, the larva is torn in half, and then is only of use for purposes of
identification. It is best to apply a small quantity of glycerine or very dilute chloro-
form water to the area where the larvae are, and then some will emerge, or so alter
their positions that they can be extracted without damage. It is very probable that
the larvae are all at the same stage, though there may be more than one species
present. As many as can be removed alive should be placed in a tube, and one or more
then dropped into boiling water in order to stretch them out and render examination
more easy. The truncated end should be examined with a hand lens, and if the
posterior stigmata are small and there are only two slits the larva is a second-stage
one. A few should now be preserved in 80 per cent: alcohol, after killing them by
dropping them into boiling water or hot 70 per cent. alcohol, and a label with details
written in pencil put in the tube, which should be well corked. It is now necessary
to rear the remaining living larvae, and this is best accomplished by placing them on
a piece of meat, or in the mouth of a dead bird or small animal. Those who have
not reared any of the larvae of the myiasis-producing Diptera may be misled by
thinking that this part of the investigation is quite easy, but I would like to warn
them that this is by no means the case; indeed it is by far the most difficult part of
such an inquiry. In theTropics great care has to be taken to prevent other blowflies,
and particularly species of Sarcophaga, from laying their eggs or depositing theirlarvae,
as the case may be, in the meat or the body as soon as it begins to decompose. I
have tried every kind of receptacle in which the meat, etc., was placed, and have even
sealed the lid with vaseline, but in every case the larvae of other blowflies, and
especially Sarcophaga, have made their way in, and then the special larvae removed
from the case of myiasis have soon disappeared. Fabre long ago suggested that a
paper bag was the best way to prevent blowflies from laying their eggs in meat ;
and taking advantage of this suggestion I placed the meat containing the larvae to be
reared in several sheets of newspaper, turning down the ends and tying it up into
a packet. No larvae can now find their way into the packet, provided that the
moisture which exudes from the decomposing meat or body does not soak right
through, and forming a wet patch under the packet, so soften the paper as to allow
small larvae to eat their way in and reach the meat. As soon as the outer part
becomes wet, the packet should be wrapped up in several fresh sheets of newspaper ;
and this may have to be repeated several times. Although large numbers of
eges may be laid under the packet, as long as the outer layers are dry the larvae
cannot make their way to the meat. When the special larvae are nearing maturity,
they crawl out into the folds of the paper and pupate, so that it ismecessary to examine
the packet for such larvae every two or three days; larvae about to pupate can
always be recognised by their yellowish-white colour and the absence of food in
their alimentary tracts. Several mature larvae should be preserved, as noted above,
the remainder allowed to pupate, and the puparia later removed and placed in tubes
with well-fitting corks. From 24 to 36 hours after the flies have hatched they should
be killed and pinned with their empty puparia ; and all dead larvae, whether first,
second or third stage, preserved in one tube containing 80 per cent. alcohol.

I have collected most of the Indian CALLIPHORINAE and their early stages by
placing dead animals out in the open, and when a female of any species came to lay
its eggs, it was carefully observed, and after a large number of eggs had been laid, in
most instances the fly was caught, pinned and identified. The mass of eggs was then
removed by cutting off a piece of tissue, so as not to injure the eggs ; a fewwere then

260 MAJOR W. S. PATTON.

placed separately in a dry tube in order to preserve the first-stage larvae as soon as they
hatched, and the remainder of the eggs were placed in the mouth of a dead bird or
animal, which was then wrapped up as explained above. Some second-stage larvae
were collected about the second or third day, the third-stage larvae later, and finally
the flies were bred out of puparia. In this way I was able to obtain the eggs, first,
second and third-stage larvae, of a large number of the commoner Indian species;
such material is extremely valuable for comparative studies.

This method of rearing the larvae of the semi-specific myiasis-producing larvae is
very satisfactory. In the case of the specific myiasis-producing species, such as
Chrysomyia bezziana or Wohlfahrtia magnifica, it is a mere waste of time attempting
to rear their larvae in decomposing animal matter. They can only be reared to
maturity in living tissues. It is therefore obviously necessary to be able to recognise
the second-stage larvae of such a species as Chrysomya bezziana—the first stage is
very seldom found, as it is a very short one. The second-stage larva of Ch. bezziana
is fortunately very characteristic, and I am not acquainted with any other species
which has a similar larva. The segmental spines are extremely well developed and
stand out as dark bands; the anterior spiracles consist of five finger-like processes ;
there is a well developed short accessory belt of spines at the sides of the abdominal
segments 1 to 7 in front of the segmental bands, and the spines on the ventral pads
consist of two separate rows. All the spines become weaker after the Sth segment,
and are poorly developed on the 8th. The posterior stigmata are situated on a well
developed plate, and consist of two rather broad straight slits directed backwards
and inwards; the tracheae give off many branches after the vestibule, so that the
Sth segment has a dark colour, owing to these showing through the integument.

If the case is one of rhinal, oral, aural or vaginal myiasis, the larvae should be
removed by spraying the parts with glycerine, or even with a little weak chloroform
water ; it may be difficult to dislodge the larvae when they are high up in the accessory
nasal sinuses, but it is nearly always possible to get a few. Medical officers in India
have always been able to send me a considerable number of larvae from such cases.
The mature larvae of Ch. bezziana when removed from the tissues should be placed
in some earth in a cigarette tin and allowed to pupate.

With the help of the above key, medical and veterinary officers should find no
difficulty in determining the genera to which the larvae belong, by simply examining
the posterior stigmata with the aid of a pocket lens magnifying 15 times. The
presence of fleshy processes on the larva points to its being either Fannia, Aphiochaeta
or such CALLIPHORINAE as Chrysomyta albiceps or vartpes, two Oriental and Australian
blowflies. If the larva is smooth, note whether the stigmatic slits are curled or
straight. If curled it belongs to the Muscinaeg, if straight, to either Calliphora,
Chrysomyia, Lucilia or Sarcophaga. With the aid of the key, and the drawings
accompanying these notes, the larva can be accurately placed in any of these genera.
If there is any doubt as to whether the chitinous ring is complete or not, shave off
the end of the 8th abdominal segment with the stigmata with a razor, and clear it
in 10 per cent. caustic potash, and mount, after dehydration, in Canada balsam on
a slide. The determination of the species should, in most cases, be left to a specialist.

All Sarcophaga larvae should be reared to maturity, and the adults hatched out
and carefully pinned, especially the males, as there is no means at present of deter-
mining the larvae of these flies. Any such material would be most valuable.

In the case of the OESTRIDAE, all of which are specific myiasis-producing flies,
success in hatching out the adults is only possible if the mature Jarvae, which have of
their own accord left their host, can be obtained. Such larvae should be placed in
some earth in a cigarette tin and should be handled as little as possible. | The mature
larvae of the OESTRINAE are usually much darker than the immature specimens;
in others the skin becomes dark yellow and harder. Mature larvae can sometimes
be obtained in places where their hosts are in the habit of lying up during the evening,

NOTES ON THE MYIASIS-PRODUCING DIPTERA. 261

for it is usually then that they leave them. The camel bot larvae always leave their
hosts in the evening, when the camels are teeding. They will be found buried in the
earth and under leaves, etc.

In the case of recently killed wild animals, such as deer, antelopes, pig, wart-hogs,
rhinoceroses, zebras, elephants, etc., if the observer has time, many larvae may be
collected from the nose, accessory sinuses, throat, nasopharynx, skin, stomach and
duodenum. The largest and the smallest larvae should be specially selected, for
in this way the nearly mature third-stage larvae, and sometimes the second stage,
will be obtained. The larvae should be placed in tubes until they can be preserved
by killing them in boiling water or alcohol. I hope those who have opportunities
while on shikar trips in Africa and India will collect any Oestrid larvae they may
find and preserve them as described above and send them to Dr. Marshall. The hosts
of many of these flies are gradually being exterminated, and with them these inter-
esting insects will disappear before we have time to learn all there is to be learnt from
their larvae.

The first-stage larvae of all oviparous OESTRIDAE can be obtained by collecting
the eggs from the hairs, skins, tusks, trunk, etc., of their respective hosts, whether
dead or alive. The hairs, or the piece of skin, should be cut off and placed in a dry
tube with a well-fitting cork, and watched every day. In a few species the larvae
hatch out of their own accord, such as those of Gasterophilus veterinus (nasalis),
Hypoderma bovis and H. lineata; others again only require moisture, when they at
once escape from the eggs, as in the case of the larvae of Cobboldia elephantis ;
and others again require friction as well as moisture, such as the larvae of Gastero-
philus intestinalis. Recently Captain Cross, C.V.D., sent me a large number of
horses’ hairs with eggs of Gasterophilus intestinalis var. bengalensis from the legs of
horses in the Punjab. The larvae could not hatch out when the eggs were only
moistened ; but as soon as they were compressed, the opercula broke away. . This
friction is applied by the tongue of the horse.

All such first-stage larvae should be dropped into hot 70 per cent. alcohol and
placed in a small tube with a good cork, which must be paraffined in order to prevent
the alcohol escaping.

In the case of the larviparous OESTRIDAE, such as the OESTRINAE, the first-stage
larvae can only be obtained by catching a female, either when depositing its larvae
at the entrance to the nose, or in the neighbourhood of its host, on the off chance
that it may contain larvae ready to be extruded. The flies should be preserved in
70 per cent. alcohol, for the larvae can always be dissected out afterwards ; and the
fly can be identified from such spirit specimens. This also applies to species of Wohl-
fahrtia and Sarcophaga, for these flies are larviparous in habit.

In concluding these notes, I would like to take this opportunity of thanking all
those who have given me material bearing on this subject. Owing to the long
distance it will of course be impossible to send me living larvae, so that the breeding
out of the flies will, in the first instance have to be carried out on the spot. But as
more material other than Indian becomes available, it will be possible to give a
complete description with a key to the larvae of the myiasis-producing Diptera, so
that a Medical or Veterinary Officer when he comes across a case may be in a position
to determine the species without having to breed out the fly ; breeding experiments
are always difficult, take up a lot of time, and, as often as not, fail. Medical and
Veterinary Officers have unique opportunities of collecting the larvae of these flies
and of studying their habits, but I believe many are deterred from doing so because
there is no easily accessible account of these interesting parasites of man and animals;
I trust these notes will in some way fill this hiatus. And lastly, I would like to point
out that comparative studies of the first, second, and third-stage larvae of these
higher Diptera cannot but lead to a better knowledge of the relationships of the adults.

me KEVISION’ OF THE MOSQUITOS OF THE PALAEARCTIC REGION,

By F. W. Epwarps.

(Published by permission of the Trustees of the British Museum.)

For some years after the intensive study of mosquitos began in tropical countries,
surprisingly little interest was taken in the European species, particularly those of
Northern Europe. Ficalbi had published his monographic revision in 1896-99,
but from then until 1914 very little further had been done ; the adults were assumed
to be more or less known, though very few of the larvae had been described. Since
1914, however, a great deal of work has been done all over Europe, bionomic as well
as systematic, and considerable advances have been made in every branch of our
knowledge of these insects.

The present paper was commenced early in 1919, with the study of a number of
large collections received at the British Museum from Italy (Mr. E. Hargreaves),
Macedonia (Capt. J. Waterston), Palestine and Mesopotamia (Capt. P. J. Barraud),
and Egypt (Major E. E. Austen). Shortly afterwards a correspondence with
Dr. Wesenberg-Lund, of Copenhagen, made it evident that there were many more
species in Northern Europe than had previously been supposed. I therefore
determined to attempt a revision of the Palaearctic mosquito fauna, and with this
end in view wrote to the Dipterists in charge at various continental museums, as
well as some private collectors, for the loan of material for determination or re-
determination. Collections were sent in response to my requests by Dr. R. Frey,
Helsingfors Museum ; Dr. E. Bergroth, Jamsa, Finland; Dr. Y. Sjéstedt, Stockholm
Museum; Dr. Wesenberg-Lund, Hillerod, Denmark; Dr. G. Enderlein, Berlin
Zoological Museum ; Dr. W. Horn, Berlin-Dahlem Entomological Museum ; Herren
L, Oldenberg and B. Lichtwardt, Berlin; M. E. Séguy, Paris Museum; Dr. H.
Zerny, Vienna Museum ; Dr. K. Kertész, Budapest Museum ; Prof. M. Bezzi, Turin ;
and odd specimens were also received from Dr. Lundbeck, Copenhagen Museum ;
and Dr. E. Martini, Hamburg. A further valuable collection was sent by Mr. S.
Yamada from Japan in 1916, and additional Japanese and Chinese collections were
made by Dr. W. A. Lamborn in 1921. To all who have helped me in this way
I wish to express due thanks, as also to other correspondents and colleagues for help
in other ways.

The publication by Dr. E. Martini of his admirable paper, ‘‘ Uber Stechmiicken,”’
in August 1920, and by Dr. Wesenberg-Lund of his equally excellent treatise on
Danish mosquitos early in 1921, have to a great extent covered the ground which
it was intended to investigate in this paper, and have also gone much further in
giving an account of the life-history of the majority of the European species. The
present paper, while not claiming to make any very considerable further contribution
to our knowledge of these species, will serve to correlate and summarise the results
of these and other workers, and by including the whole Mediterranean region give a
rather more comprehensive view of the subject.

The geographical limits adopted here are those most usually given to the Palae-
arctic region, 2.¢., Europe, North Africa, as far south as the tropic; the Atlantic
Islands ; Asia Minor; North Arabia, including the head of the Persian Gulf ; North

264 F. W. EDWARDS.
Asia as far as the Himalayas; North China; Japan. Mosquitos have been more
or less carefully collected over practically the whole of this area, the least known
portions being perhaps North Russia and North and Central Siberia. Spain also
has not yet been thoroughly worked. Iceland should be included in the area, but
no mosquitos have yet been recorded or received from that country.

The study of the various collections enumerated above indicates that the mosquito
fauna is fairly homogeneous over the greater part of the area, but there are two parts
which, so far as this group of animals is concerned, would seem to be better classed
in the Oriental region. These are the area immediately round the head of the Persian
Gulf, and the southern islands of Japan, at least as far north at Tokio. In the former
area there is a very large, perhaps a predominating admixture of Oriental forms,
while round Tokio the fauna appears to be of an almost purely Oriental type. If
these two areas had been excluded from consideration, the number of species dealt
with would have been much less; but it was thought that the usefulness of this
paper might be increased by keeping to the wider limits.

The total number of species dealt with is as follows :—

Anopheles, 19. Orthopodomyra, 1.
Uranotaenia, 2. Taenorhynchus, 1.
Rachionotomyia, 1. Aédes, 38.
Megarhinus, 3. Armigeres, 1.
Theobaldia, 7. Lutzia, 1.

Culex, 20.

The faunistic relationship of these species may be analysed as follows :—

Intrusions from the Oriental Region. Intrusions from the Ethiopian Region.
In the West. In the East. In Algeria, etc. pees and
alestine.
A. stephensi. An. lindesayi, Arm. obturbans. An, costalis. A, mauritianus.
C.tritaeniorhynchus., U.bimaculata. L. vorax. Ae, vittatus. A. pharoensis.
C. vishnui. R. bambusa. — C. hayashi. C. quasigelidus,
C. fatigans. M.towadensis. C, bitaeniorhynchus, |y——————<“—s~Sts—tS CC... Laveen.
A, togoi, C. sinensis, In Mesopotamia. |C.tvitaeniorhynchus.
A. japonicus. C, tritaeniorhynchus.
A. koveicus, C. vishnut.
A. niveus. C. fatigans. A. rhodesiensis,
A. albopictus. -

The remaining 70 species may be said to comprise the true Palaearctic fauna
(though a number of them spread into adjoining regions). As might be expected,
there are considerable differences between the North European and the Mediterranean
faunas. The former shows very strong affinities with the Nearctic fauna, and a con-
siderable number of the species seem to be common to both Europe and North
America, while others have obviously representative forms in the two regions. The
species which I consider either identical or only varietally distinct are: Anopheles
maculipennis, Theobaldia alaskaensis, Aédes dorsalis, A. lutescens, A. excrucians,
A. alpinus, A. cataphylla, A. diantaeus, A. sticticus, A. punctor, A. communis,
A. intrudens, A. pullatus, A.vexans, A. cinereus, Culex apicalis, C. pipiens. Nearly
all these species are already known to occur throughout Europe and Siberia, and the
two faunas may therefore be assumed to have mixed at a recent date by way of
Eastern Siberia and Alaska. In one or two cases (e.g., Anopheles maculipennis)
there seems to be definite evidence that the European fauna is more closely allied
to that of Western than that of Eastern North America.

MOSQUITOS OF THE PALAEARCTIC REGION. 265

The following may be considered representative species :—
Europe. North America.
Anopheles plumbeus. A. barbert.
Theobaldia annulata. T. maccrackenae.
T. glaphyroptera. T. impatiens.
T. morsitans. T. dyari.

Orthopodomyia pulchripalpis. O. signifer.
Taeniorhynchus richiardi. T. perturbans.
Aédes semicantans. A. stimulans.
A. rusticus. A. trichurus.
A. geniculatus. A. triseriatus.

It is of special interest to note that none of the European species in this list
are at present known from Asia.

As might be anticipated, few, if any, of the purely southern species have any
clearly recognisable North American representatives, but some of them have wide
extensions of their range into the Ethiopian and Oriental regions. The most note-
worthy of such species are Theobaldia longiareolata and Culex tipuliformis.

In the more northerly parts of the region the dominant group is the subgenus
Ochlerotatus of Aédes; the species of Anopheles which occur are all of the typical
subgenus. Further south Ochlerotatus rapidly disappears, and begins to be replaced
in part by species of Culex, and in part by other subgenera of Aédes, while the
Anophelines of the Myzomyra group become numerous.

I have endeavoured in this paper to revise the generic classification of the CULICIDAE
on a sounder basis than has been used hitherto, by defining the genera primarily
on characters which have no relation to sex. In previous papers I have tried to
minimise the use of secondary sexual characters, as well as of the merely superficial
characters of the scales, but found myself in some cases compelled to fall back upon
these, or else on the male hypopygium. A closer study of the insects now enables
me to point out what appear to be constant and important distinctions between
most of the genera which are applicable equally to both sexes without being so
artificial as the scale characters have been proved to be. Fortunately no changes
are involved in our conception of the limits of the genera; these had been already
soundly established by the study of the larvae, and the new characters adduced
only confirm most of the conclusions already arrived at by a study of the early stages.

Several of the new distinctions employed are to be found in the thoracic chaetotaxy,
and to explain these clearly I give explanatory figures of the pleural bristles in
Theobaldia annulata and Aédes geniculatus. The most important bristles for taxonomic
purposes I find to be those occurring on the area in front of the prothoracic spiracle.
This area is almost entirely occupied by the proepimeron, but there is a small, more
or less triangular area immediately adjoining the spiracle which is separated from
the proepimeron by a well-marked ridge. The bristles which are found on this small
area I speak of here as the spiracular bristles. When present they project backwards,
covering and protecting the spiracle. When the spiracular bristles are absent,
their function is often assumed by the proepimeral bristles, which are usually situated
in a row near the posterior margin of the proepimeron ; it will sometimes be necessary
to look closely in order to ascertain whether the bristles present are spiracular or
proepimeral, or perhaps both ; their position relatively to the above-mentioned ridge
is the deciding factor. The two groups are obviously not homologous.

The other groups of bristles on the pleurae are apparently of less importance, and
are for the most part more hair-like. They are as follows :—Pvonotal, on the pro-
thoracic lobes. Prosternal, on the prosternal lobes; a group which Christophers
considers important in the ANOPHELINI, but which seems to be fairly constant through-
out the Curicini. Post-spiracular, a distinct group of some importance situated
a little behind the prothoracic spiracle and below the margin of the mesonotum.

266 F. W. EDWARDS.

Pre-alar, a clump situated on the pre-alar prominence ; always present, but variable
innumber. Sternopleural, the row which is nearly always present, extending vertically
across the mesepisternum and mesosternum, almost in a line with the row on the

Fig. 1. Side view of thorax of (A) Theobaldia annulata and (B) Aédes

geniculatus, to show arrangement of pleural bristles. (Scales omitted.)

Bristles: p.n., pronotal; p.s., prosternal; p.e., proepimeral; sp., spiracular ;

p. sp., post-spiracular; p.a., pre-alar; wu. mz.e., upper mesepimeral ;
l.m.e., lower mesepimeral, st, p., sternopleural.

middle coxae. Mesefimeral, two groups whose position is indicated by the name ;
the upper mesepimeral bristles form a distinct clump or tuft which is always present ;
the lower mesepimeral are variable in number and have been used to distinguish the

genera Culex and Lutzia.

MOSQUITOS OF THE PALAEARCTIC REGION. 267

I have studied also the mesonotal bristles, but although these vary greatly in
number and development, I have not succeeded in discovering any differences which
are of more than specific value. The same applies to the bristles of the legs, but those
of the head sometimes offer useful generic characters. There are two great advantages
which bristle characters have over scale characters. Firstly, they are obviously
of much greater phylogenetic importance. Secondly, even when they are rubbed
off they always leave a recognisable scar. A denuded specimen may even be more
easy to examine than a perfect one, as the scales are sometimes liable to obscure the
bases of the bristles.

In this paper I believe I have mentioned every name proposed for a mosquito
taken within the region dealt with, but I have not as a rule given additional synonyms,
nor have I attempted to unravel all the confusion caused by mis-identification, but
have tried merely to give clear definitions of the specific concepts. A large number
of the old descriptions are unrecognisable, and the types of many (some of Meigen’s
and all of Robineau-Desvoidy’s) have apparently ceased to exist. Since, however,
it is highly probable that the old names all apply to species which are known at the
present day, I have endeavoured to form an opinion as to which species was most
likely intended by the describer, rather than give a separate list of indeterminable
species. I have not included Robineau-Desvoidy’s Culex flavovirens and C. viridis
in the synonymy, because I consider it obvious that they were CHIRONOMIDAE.
Linnaeus’ Culex vulgaris may very likely have been a species of Aédes, but has
been considered by Dyar and Knab to be a Simulium, it is therefore omitted.
Anopheles sacharovii, A. pseudopictus var. flerowi, Portch., and A. superpictus var.
vassilievi, Portch., referred to by Russian writers (see Review of Applied Entomology,
B, ii, p. 108, and iii, p. 196), do not appear to have been described; at any rate
I have been unable to trace the descriptions. For references to the original
publications of Culex annulatus, de Fourcroy, C. miveus, Eichwald, and C. pallies,
Waltl, I am indebted to Mr. C. Davies Sherborn, who has kindly allowed me access
to his manuscript.

Tribe ANOPHELINI.
Genus Anopheles, Mg.

As the careful and detailed researches of Christophers have shown, there are
two main sections of the genus Anopheles, differing in small but quite easily definable
characters, both in the adult and in the larva, and undoubtedly to be regarded as
representing a very early separation of the genus into two distinct stocks. This
separation, according to Christophers, probably took place at least as early as the
Cretaceous. It is both desirable and convenient to recognise this important conclusion
in our nomenclature, and I therefore accept the divisions proposed by Christophers
(Ind. Journ. Med. Res. iii, p. 383, 1915) as subgenera. I do not consider that the
differences are sufficiently important or sharply defined to warrant the full generic
separation of the two old-world groups, nor do I consider it possible to subdivide
either of these groups in a satisfactory manner, since the connections between their
component species are so intricate and the intergradations so complete. As
Christophers has pointed out, the subgenus Anopheles shows much more real diversity
among its members than the subgenus Myzomyia, but when the species of the whole
world are taken into consideration it is impossible to recognise clearly-defined
divisions.

The eggs of different species of Anopheles show remarkable variations in the
structure of the air-floats, while these appear to be constant for each species ; in
some instances, indeed, this is so markedly the case that the egg-structure (when
known) affords the readiest means of distinguishing closely allied forms. I have
therefore considered it worth while to collect into one block all the published figures
of the eggs of Palaearctic Anopheles. These, with the two or three new ones which

268 F. W. EDWARDS.

are added, comprise all the species dealt with in this paper, except A. mauritianas
and A. sergenti. It will be noted that the species in which the air-floats have under-
gone great reduction are A. plumbeus, A. elutus, A. multicolor and A. turkhudt.
Dr. G. A. K. Marshall suggests that this reduction may be connected with the habit

ead

eee!
SF

SQW

so jee
S

Fig. 1.— Eggs of Palaearctic species of Anopheles. a, A. plumbeus, Steph. (original) ;

b, A. algeriensis, Theo. (after Sergent) ; c, A. bifurcatus, L. (after Martini) ; d. A. maculipennis,

Mg. (original) ; e, A. elutus, sp. n. (original) ; f, A. hyvvcanus (Pall.) (original) ; g, A. pulcherrimus

Theo. (after Stephens and Christophers) ; h, A. stephensi, Theo. (after Stephens and Christophers) ;

i, A. costalis, Theo. (after Patton); 7, 4. rhodesiensis, Theo. (after Patton); k, A. multicolor,

Camb. (after Foley) ; /, 4. turkhudi, Liston (after Stephens and Christophers) ; m, A. hispaniola,
Theo. (after Sergent).

of breeding in water of high specific gravity (tanninised water in the case of A.
plumbeus, saline water in the case of A. multicolor), in which large floats would perhaps
not be necessary in order to keep the eggs on the surface during incubation. It is
uncertain whether this explanation will apply to A. elutus and A. turkhudt.

Subgenus Anopheles, Christophers.

Ind. Journ. Med. Res. iii, p. 383 (1915).

Adult.—Cross-veins and bases of forks of wing-veins with dark scales. Costa
largely dark (in Palaearctic species), with at most two pale spots, apart from a pale
area in the apical fringe. Male hypopygium with two (rarely one or three) strong
spines at the bases of the side-pieces, one or both of which are borne on a strong
tubercle. Prosternal hairs usually numerous.

MOSQUITOS OF THE PALAEARCTIC REGION. 269

Larva.—Shaft of antenna with a branched hair (except in A. plumbeus). Palmate
hairs lanceolate, without long terminal filament. Internal clypeal hairs generally
close together.

The members of this subgenus exhibit considerable diversity, and some obviously
fall together into groups, but when the species of the whole world are considered it
does not seem profitable or even possible to recognise distinct subgenera. The nine
Palaearctic species at present known may be distinguished by the following tables :—

10.

. Base of side-piece with only one strong spine us .. algeriensis, Theo.

Adults.

Wing-scales all dark; front femora cylindrical ; female palpi slender,
the scales appressed ( Anopheles) ne oe i a
Wings with many pale scales; front femora thickened at the base ;
female palpi and proboscis (except in <A. lindesayt) shagelly scaled
towards the base (Myzorhynchus) .. : oe TO
Wing-scales uniform ; base of upper fork- call neta nearer wing-
base than that of the lower ahs 3
Wings normally with spots of darker, more “densely ag. geregated ‘scales :
fork- cells somewhat shorter, their bases practically level a rie as

_ White frontal tuft absent ; mesonotum unicolorous, with dark hair

algeriensis, Theo.
White frontal tuft present ; mesonotum darkened at sides, with some

pale hair and scales .. ies Sir) Bae
. Smaller, blacker species ; a distinct nate of wane eats on Mee ont i meso-
notum os ; .. plumbeus, Hal.
Larger, browner species : " scales on front margin of mesonotum narrower
and not pure white = ag e bifurcatus, L.
. Mesonotum unicolorous ; wing-fringe all Haste : of elutus, Sp. N.

Mesonotum darker at sides ; fringe “usually pale at tip of wing
maculipennis, Mg.

. Costa with only one pale spot (at the tip) ; hind femora with a conspicuous

white pre-apical ring; front femora only slightly enlarged at the base
lindesayt, Giles

Costa with two distinct pale spots (normally) ; hind femora without pre-

apical white ring ; enlargement at base of front femora conspicuous .. 7

. A white dot at base of costa, anda second nearit .. .. punctibasis, sp. Nn.
These dots absent se ve We hye ev Be Mahe)

. Fourth and fifth hind tarsal fone all nite Sy . .. mauritianus, Grp.
Fifth hind tarsal joint dark... is of a cae eo)

. Fourth hind tarsal joint dark sents at aie ap oF 10

Fourth hind tarsal joint all white or whitish, Ayrcanus var. “pseudopictus, Grassi
Wing-markings usually more pee defined ; basal half of costa with

Gare scales only : te sae hyrcanus, Pall.
Wing-markings more blurred ; “basal “half” of costa with some pale
scales on its inner edge... sre hyrcanus var. mesopotamiae, Chr.

Male Hypopyguun.

Base of side-piece with more than one strong Spimes-.t 2

Base of side- -piece with three spines, two of them br scned tones be

Base of side-piece with two spines, both simple ae ae i Pe eo.

Spines of claspette approximated, but not fused... oe che ea ax
5

Spines of claspette fused into a spatulate process

(4183) U

270 F. W. EDWARDS.

4. Aedoeagus short and broad, without leaflets .. i .. plumbeus, Hal.
Aedoeagus long and narrow, with leaflets at the tip
maculipenms, Mg. ; elutus, sp. n.
5. Spatulate process long and narrow ; ninth tergite without definite pro-
cesses ; long hair at middle of side-piece moderately developed
lindesayi, Giles
Spatulate process shorter and broader ; ninth tergite with distinct pro-
cesses ; long hair at middle of side-piece stouter, almost spine-like .. 6
6. Processes of ninth tergite short and broad .. es punctibasis, sp. n.
Processes of ninth tergite elongate .. /yrcanus, Pall.; mauritianus, Grp.

Larvae.
1. *\ row of six large plumose hairs across middle of head... wil at ee
These hairs vestigial, simple .. os , .. plumbeus, Hal.
2. Outer anterior clypeal hair thickly branched, Ribiire A a os FES
Outer anterior clypeal hair simple or slightly branched... bifurcatus, L.
3. Innermost shoulder-hair branched from base or nearly .. mauritianus, Grp.
Innermost shoulder-hair branched only at tip, if at all “G sis ee
4. Palmate hairs on abdominal segments i—vii (always ?) -. hyrcanus, Pall.

Palmate hairs on abdominal segments 1ii—vii only
elutus, sp. n.; maculipenms, Mg.
The larvae of A. algeriensis and A. lindesayi are insufficiently described for
inclusion in the above table; that of A. punctibasis is not yet described.

1. Anopheles (Anopheles) algeriensis, Theo. (fig. 2, b).
Anopheles algeriensis, Theobald, Mon. Cul. iii, p. 21 (1903).
Anopheles lukisi, Christophers, Ind. J. Med. Res. iv, p. 120 (1916).

Fasily distinguished by the unspotted wings and the absence of a distinct tuft
of long white scales on the front of the head. The mesonotum is uniformly dull light
brown, and clothed with dark hair only, the small yellowish hairs or hair-like scales
which are found in A. bifurcatus being absent in this species. In the female the second
segment of the palpi is considerably longer than the first or the third and fourth
together.

The male hypopygium, which has not as yet been described, is very distinct.
The processes of the ninth tergite (ventral processes of Christophers) are very short,
less than half as long as the breadth of the tergite. The side-pieces have a strongly
differentiated bristle on the inner side near the tip, as in A. bifurcatus ; there is a
single very strong basal spine, situated on a large tubercle, with its tip bent; the
outer basal spine is absent or represented only by one or two scarcely differentiated
and inconstant bristles. The claspettes are trilobed, the first lobe bearing two or
three strong, pointed, closely approximated spines, the second several short hairs,
the third (innermost) three nearly equal, moderately strong bristles. The aedoeagus
bears two or three pairs of long delicate leaflets at its tip.

The larva is unknown. Sergent, indeed, gives a short, incomplete description
and some figures of a larva which he supposes to be A. algertensis, but he was unaware
of the real distinctions between A. algeriensis and A. bifurcatus, and his statements
are insufficient to enable us to decide which of these species he had before him, or
whether he had both. Sergent’s figure of the egg is reproduced in fig. 2, 6 ; it is not
improbable, however, that this really represents the egg of A. bifurcatus.

The type of A. lukisi shows faint traces of pale banding on the hind tarsi; in
the Mediterranean specimens the tarsi are entirely dark, but there appear to be no
other differences. The colour of the mesonotum varies from yellowish to rather
dark brown.

MOSQUITOS OF THE PALAEARCTIC REGION. ATM |

The species has a rather close resemblance to the Oriental A. aitkent, and, as
suggested by Barraud, it is probable that the specimens of A. aitkent recorded from
Palestine were really A. algeriensis. A. aitkeni is certainly a distinct species, since
the upright scales of the head are very much narrower than in A. algertensts, and
the side-pieces of the hypopygium have two distinct basal spines.

Distribution.—Algeria (Sergent) ; Italy (Hargreaves) ; Macedonia (Waterston) ;
Palestine (Austen, Barraud) ; Egypt (Austen) ; Mesopotamia (Christophers).

2. Anopheles (Anopheles) bifurcatus, L. (fig. 2, c).

Culex bifurcatus, Linnaeus, Syst. Nat. Ed. x, p. 603 (1758).

Culex claviger, Meigen, Klass. u. Beschr. i, p. 4 (1804).

Culex trifurcatus, Fabricius, Ent. Syst. iv, p. 401 (1794).

Anopheles villosus, Robineau-Desvoidy, Mém. Soc. d’Hist. Nat. Paris, i, p. 411

(1827).

Anopheles grisescens, Stephens, Zool. Journ. xii (1828).

Anopheles antennatus, Becker, Mitt. Zool. Mus. Berlin, ii, p. 68 (1903).

This species varies a good deal in size and colour ; small dark specimens might

be mistaken for A. pluwmbeus and small light ones for A. algeriensis. Apart from the
male hypopygial characters, which are absolutely diagnostic, A. bifurcatus may
readily be separated from A. plumbeus and A. algeriensis by the other characters
mentioned in the key.
’ The larva, which has been fully described by Lang, occurs generally in clean
water with some weed, in rather shady situations ; in the Mediterranean region it
is frequently found in wells. The winter is passed in the larval stage, the adults
dying off in the late autumn and the first brood appearing again in early spring.

A. bifurcatus is less domestic than A. maculipennis and does not so readily
enter houses, but bites freely in the open.

Distribution :—Throughout Europe, North Africa (except desert regions) and
Asia Minor. Turkestan (Vassiliev). Not found as yet furthereast. Apparently rarer
in the far north.

South European specimens are commonly smaller than those from Central and
North Europe, but are not otherwise distinguishable.

3. Anopheles (Anopheles) plumbeus, Stph. (fig. 2, a).

Anopheles plumbeus, Stephens, Zool. Journ. iii, p. 503 (1828).
Anopheles nigripes, Staeger, Kroyer’s Nat. Tidschr. ii, p. 552 (1839).

The general black colour, densely and uniformly scaled wings, white scales on
front of mesonotum, conspicuously hoary stripe down centre of mesonotum, ‘and
conspicuous white frontal tuft, will suffice to distinguish this species from those
with which it might be confused.

The larva and egg have been described by Lang, Eysell, and Carter and Blacklock.
A. plumbeus is the only species in the Palaearctic fauna which is known to breed
in rot-holes in trees, and the very striking characteristics of the larva all seem to be
correlated with this habit. As has already been pointed out by Christophers, tree-
hole larvae of different genera of mosquitos commonly show three forms of modifi-
cation : (1) a reduction of the hairs on the head, and especially of the small bristles
on the antennae; (2) an increase and enlargement of the hairs on the thorax and
abdomen ; and (3) a development of chitinous structures on the last few abdominal
segments. All these points are well illustrated by A. plumbeus. The tree-holes
in which these larvae are found are frequently (perhaps usually) dark, and it may be
supposed that the sense of sight would be of little value to the inhabitants of such
places, while that of touch would attain a greatly enhanced importance, since the

(4183) u2

O72 F. W. EDWARDS.

mosquito larvae have a number of predaceous enemies living with them. Possibly
this may help to explain the development of hairs on the soft parts of the body,
though their reduction on the head is not so easy to account for. The development
of additional chitinous plates may be merely a chemico-physical reaction to the excess
of tannin in the water.

Whatever may be the use of these larval modifications, it seems certain that they
are due to environmental conditions (since they are shown by species in totally
unrelated genera), and therefore the genus Coelodiazesis, founded solely on these
characters, cannot be satisfactorily maintained.

Distribution.—Throughout Europe, wherever there are many deciduous trees,
in which rot-holes can form. Not yet recorded with certainty from North Africa
or Asia, except one from Katmia, Cilicia (Lt.-Col. Lelean, recorded by Christophers).

The Himalayan species A. bavianensis, James, has been referred to by Christophers
as identical with A. plumbeus, but there are small differences: in A. bavianensis
the white scales on the mesonotum extend further (almost half-way from the front)
and the femora and tibiae are conspicuously white at the tips, whereas in A. plumbeus
they have scarcely a trace of white. As there are also small distinctions between the
larvae, it will probably be best to regard the two as distinct species, occupying
separate geographical areas in the Palaearctic and Oriental regions. The type has
also a North American representative in A. barberi, Coq., which is distinct from both
the old-world forms, though closely resembling them.

4. Anopheles (Anopheles) maculipennis, Mg. (fig. 2, d).

Anopheles maculipennis, Meigen, Syst. Beschr. i, p. 11 (1818).

Culex claviger, Fabricius (nec Meigen), Syst. Antl. p. 35 (1805).

Anopheles occidentalis, Dyar & Knab, Proc. Biol. Soc. Wash. xix, p. 159 (1906) ;
Howard, Dyar & Knab, Mosq. N. & C. Amer. iv, p. 1026 (1917).

Anopheles lewisit, Ludlow, Psyche, xxvii, p. 74 (1920).

Anopheles selengensis, Ludlow, Psyche, xxvii, p. 77 (1920).

This could not easily be confused with any other Palaearctic species except
A. elutus; from this it differs in several small details of coloration, as indicated
in the key. The pale area in the fringe at the tip of the wing is diagnostic of A.
maculipennis, when it is present, but can only be seen in perfect specimens, the fringe
at the wing tip being very easily denuded ; moreover, the fringe is entirely dark in
some individuals, and as the wing-spots also vary somewhat in intensity, the distinction
between the adults of these two species is not very clearly marked. Perhaps the
best distinction between the two species is in the colour of the scutum, that of
A. maculipennis having a broad brown or blackish-brown stripe on each side, as in
A. bifurcatus and A. plumbeus.

According to Wesenberg-Lund the Southern European race of A. maculipennis
is smaller than the Northern. The difference, if it exists, is very slight, and is only
to be found in the average measurement, since many Mediterranean specimens which
I have seen are quite large. Dyar (in correspondence) also maintains that there is
a minute difference in the hypopygia between specimens from France and Siberia
and those from Hungary. I am unable to confirm this, and consider that the small
amount of variation which does occur is individual only. Mounts of male hypopygia
of specimens from Britain, Macedonia and Constantinople show no difference whatever.

In my opinion the Western North American A. occidentalis is specifically identical
with A. maculipennis. I can discern no difference in the larvae; the adults are
alike in all external features, and the only demonstrable difference in the male
hypopygia is that the two outer spines on the claspette are both pointed in
A. occidentalis, while one or both of them are blunt-ended in A. maculipennis.
Such a distinction seems to me inadequate even for varietal separation. The egg of
A. occidentalis is as yet undescribed, and may differ from that of A. maculipennis, but

MOSQUITOS OF THE PALAEARCTIC REGION. 273

I do not anticipate that such will be found to be the case. In any event A. occidentalis
is much closer to A. maculipennis than is the Eastern North American A. quadri-
maculatus, which differs in larva, hypopygium, and adult coloration. This is a fact
of considerable interest, and is in line with what has been found in the genus A édes.

The early stages have been described in detail by Nuttall and Shipley, Lang and
others. The larvae are found most commonly in open, weedy water in low-lying
districts, sometimes in brackish water near the coast.

Distribution.—Throughout Europe, from the Arctic to the Mediterranean ; North-
West Africa (except desert regions) ; and across Siberia to North-Western America ;
also recorded by Christophers from Upper Mesopotamia.

5. Anopheles (Anopheles) elutus, sp. n. (fig. 2, e).

Anopheles sp., Christophers, Ind. J. Med. Res. vii, p. 711 (1920).
Anopheles maculipennis var., Barraud, Bull. Ent. Res. xi, p. 389 (1921).

Differs from A. maculipennis as follows: No trace of a pale spot in the fringe
at the tip of the wing, the fringe being uniformly dark. Dark spots at the bases
of the fork-cells and at.cross-veins very poorly developed, often hardly perceptible,
especially in the male. White frontal tuft small, rather inconspicuous. General
coloration of the body rather lighter than in A. maculipennis, the sides of the scutum
not any darker than the middle. Male hypopygium apparently identical with that
of A. maculipennis in structure, but the ninth tergite less strongly chitinised
relatively to the rest of the organs. Egg without lateral float-cells, evenly fringed
all round as in A. plumbeus, but resembling A. maculipennis in its more elongate
shape.

Larva apparently identical with that of A. maculipennis.

Type, a male in the British Museum reared from larva taken in marsh at
Kishon, Palestine (Capt. P. J. Barraud).

The credit for the recognition of this species belongs to Major Christophers, who
was the first to obtain the eggs. These are so different from those of A. maculipennis
that it is impossible to regard the two forms as varieties of one species, notwithstanding
the identity of the larvae and the close similarity between the adults. Major
Christophers’ observation has been amply confirmed by Capt. Barraud, who reared
the species in Palestine, and presented material of all stages to the British Museum.

Distribution.—Mesopotamia (Christophers); Palestine (Barraud); Syria
(Barraud) ; Macedonia (Waterston) ; Cyprus (Dr. G. A. Williamson) ; Transcaspia
(Amudaria, C. Ahnger, in coll. Helsingfors Mus.) ; West Caspian (Adzikabul, near
Baku, per Col. Wenyon) ; Steiermark (Admont, Sérobl.; in coll. L. Oldenberg) ; W.
Persia (Ountul, H. &. Shorit). .

In Palestine, Lower Mesopotamia and Transcaspia this species entirely replaces
A. maculipennis, but in Macedonia the two occur together. A. elutus appears
to be absent from Central and Northern Europe.

6. Anopheles (Anopheles) lindesayi, Giles.

Anopheles lindesait, Giles, Gnats, p. 166 (1900).

A very well-marked species, with no close ally, unless the Japanese and Formosan
forms are regarded as distinct. In several respects it is intermediate between the
Anopheles and Myzorhynchus groups ; the transition is perhaps made more complete
by A. gigas.

A. lindesayi is usually found at high altitudes and breeds in mountain streams.

According to Christophers’ description and figure the ninth tergite of the male
hypopygium is broad and shield-like, but in the Japanese male I have examined it
has the usual narrow form, a narrow, curved strip, the ends being pushed out but

274 F. W. EDWARDS.

not forming definite processes. This may perhaps indicate a specific difference from
the Indian form, though I cannot find any other characters in the adult to support
such a conclusion.
This is possibly the so-called new species recently recorded (but not named)
from Japan (see Tropical Diseases Bulletin, xvi, 1920, p. 106).
Distribution—Mountains of India; Formosa (Kotdzuwmi); Japan (Nagasaki,
Lamborn).

7. Anopheles (Anopheles) punctibasis, sp. n.

Closely related to A. hyrcanus (Pall.), but differs in the following particulars :—
Average size somewhat larger ($4-5mm., 95-5mm. in length of wing); wings
more distinctly mottled in naked-eye appearance than in the most brightly-marked
A.hyrcanus. Antennae slightly but distinctly more slender than those of A. hyrcanus
except for the first joint, which is distinctly stouter. Scales of antennae light brown
and confined to the torus and the first two flagellar joints (in A. hyrcanus they are
white and extend to the fourth or fifth flagellar joints). Palpi normally entirely
black-scaled. All the tibiae, the first three joints of the front and middle tarsi, and
the first four joints of the hind tarsi, narrowly but distinctly whitish-ochreous at the
base as well as at the tip. A white dot at the extreme base of the costa, and a second
immediately beyond the humeral cross-vein. Dark and light areas on veins sharply
defined, no intermixture of light and dark scales except to a very slight extent on
the third vein. Processes of ninth tergite of male hypopygium very short and broad,
almost semicircular in form (hypopygium otherwise much like that of A. hyrcanus).

Distribution—Japan : Nagasaki, 8-19.v.1921, 2 ¢ (including type), 5 Q reared
from pupae found in a muddy shaded pool in company with Culex hayashi ;
Yokohama, vi.1921, 1 9 reared from larva (Dr. W. A. Lamborn).

The sum of the characters enumerated above seems to be quite sufficient to
distinguish this form specifically from A. hyrcanus, though the close relationship
is evident. The black-scaled palpi at first sight afford an easy means of distinction,
but the Yokohama specimen has narrow white rings at the articulations and a distinct
white tip to the last joint.

8. Anopheles (Anopheles) hyrcanus, Pallas (fig. 2, f).

Culex hyrcanus, Pallas, Reise durch versch. Prov. d. Russ. Reichs. i, p. 475 (1771).

Anopheles sinensis, Wiedemann, Aussereurop. zweifl. Ins. i, p. 547 (1828).

Anopheles pictus, Loew, Dipt. Beitr. i, p. 4 (1845).

Anopheles pseudopictus, Grassi, Atti R. Acc. Lincei, Rendic. viii, 1, p. 102 (1899).

Anopheles sinensis var. mesopotamiae, Christophers, Ind. i Med. Res. iii, p. 196
(1916).

Easily distinguished from all other species in the Palaearctic fauna, except
A. punctibasis and A. mauritianus, by the distinctly swollen front femora. This
character was pointed out by Loew in his description of A. pictus, and together with
the shaggily-scaled female palpi forms the best distinguishing mark of the Myzorhynchus
group, the character of the ventral scale-tuft, on which the group was originally
founded, being quite unreliable. The relation with the Anopheles group (in the strict
sense) is however, obviously close; there is no definable difference in the larvae
or hypopygia, and the colour differences are bridged by such species as A. gigas.
I therefore do not admit Myzorhynchus as a distinct subgenus.

A. hyrcanus is a variable species in many respects, and shows a strong tendency
to the production of local races. In the Oriental region some of these may be said
to have reached specific differentiation (e.g., A. sepavatus, Leic.), but this can hardly
be said of the Palaearctic forms, at least with our present knowledge of them.

MOSQUITOS OF THE PALAEARCTIC REGION. 279

Christophers seems inclined to distinguish the var. mesopotamiae on account of a
slight difference in the shape of the processes of the ninth tergite of the male hypo-
pygium, but this distinction is no more constant than the colour characters which he
adduces to separate mesopotamiae from hyrcanus. The form inhabiting Japan
and Eastern Siberia has the wing-markings blurred somewhat as in the var. meso-
potamiae, but the general coloration is much darker. The var. pseudopictus is at first
sight sharply distinguished byits pale fourth hind tarsaljoint (one Macedonian example
has the third joint also broadly white at the tip), but the paleness in some specimens
is only visible in certain lights, and there appears to be no other distinction between
the two forms. The fifth hind tarsal joint in psewdopictus remains dark, but in
Swellengrebel’s avgyvropus from Sumatra both the fourth and the fifth joints are
white ; this condition seems to have been developed independently in argyropus
and mauritianus, and, if so, argyropus should perhaps be regarded as an extreme form
of hyrcanus rather than as an Oriental form of mauritianus.

A. hyrcanus is found chiefly in large marshes, especially near the sea. The
larva has been described by Joyeux.

Distribution North Mediterranean coast from the Rhone delta to the Levant, and
across Central Asia from the Black Sea coasts to Japan; also (in a darker variety)
throughout the Oriental region. The var. mesopotamiae appears to be confined to
the region at the head of the Persian Gulf; the var. pseudopictus has been found
in Italy (Grassi) ; Macedonia (Waterston) ; Danube delta (Leon) ; Transcaspia (Tedjen,
C. Ahnger).

9. Anopheles mauritianus, Grp.

Anopheles mauritianus, Grandpré, Planters’ Gazette Press (1900).
Anopheles paludis, Theobald, Royal Soc., Rept. Malaria Com. p. 75 (6th July,
1900).

This is the African representative of A. Ayrcanus, from which it differs almost
solely in the generally blacker colour and the white tip of the hind tarsus. The
range‘of variation of the two forms is, however, quite distinct, and it is no doubt
justifiable to regard them as separate species. Christophers distinguishes the
hypopygium from that of A. hyrcanus by the absence of leaflets on the aedoeagus,
but either he was mistaken in his observation or the species is variable in this respect,
since delicate leaflets are certainly present in those which I have examined.

Like its ally A. hyrcanus, this species lives chiefly in large swamps, especially
near coasts, and is a poor carrier of malaria.

Distribution —Tropical Africa, extending into the Palaearctic region only in the
south-eastern Mediterranean, where it occurs in the Nile delta and in the marshes
on the Palestine coast (Barraud).

Subgenus Myzomyia (Blanchard), Christophers.
Ind. Journ. Med. Res. ii, p. 383 (1915).
Adult.—Cross-veins and bases of forks of wing-veins with light scales (except
in A. rhodesiensis). Costa with four or more pale spots. Male hypopygium with
a group of several (4-6) stiff bristles at the base of each side-piece, none of which are

borne on tubercles or otherwise differentiated. Prosternal hairs nearly always
reduced.

Larva.—Shaft of antenna without a branched hair. Leaflets of palmate hairs
generally with long terminal filament. Internal clypeal hairs rather wide apart.

The members of this subgenus, though showing a great total range of ornamen-
tational characters, are really all much more closely allied than the species of the
subgenus Anopheles. The male hypopygia are almost identical throughout the
subgenus, the minute distinguishing characters given by Christophers being in some

276

F. W.. EDWARDS.

cases of very uncertain value; the species most easily recognisable as regards
hypopygial structure are those (pulcherrimus, multicolor) which have no leaflets
on the aedoeagus, but it is impossible to consider a common lack of these structures
as indicating Telationship. The larvae also are extremely similar and difficult to

distinguish.
Adults.

1. Abdomen densely scaly ; the scales forming lateral tufts on each seg-
ment ; last hind tarsal joint white; female palpi shaggily scaled .. 2

Abdomen without lateral scale-tufts; last hind tarsal joint dark ;
female palpi with most of the scales appressed... Os ee ‘aca
2. Fourth hind tarsal joint all white ws a 3 pulcherrimus, Theo.
Fourth hind tarsal joint mostly dark .. or AS ... pharoensis, Theo.
3. Femora and tibiae pale-spotted sr ue tA - - og EL
Femora and tibiae not pale-spotted ; abdomen without scales... See
4. Abdomen scaly almost to the base .. Be os .. stephensi, Liston.
Abdomen without scales, except on cerci se: ie ~ costalis, Theo.
5. Pale wing-markings confined to costa and first vein .. . .rhodesiensis, Theo.
Pale markings on all wing-veins - “it eng si ty. oy AO
6. Tip of last palpal joint white-scaled .. sha a ss oe Sed.
Tip of last palpal joint black-scaled .. ae ae are 3)

7. Thorax with hairs neue last -two white rings at jeutaie palpi quite
narrow ais sergentt, Theo.

Thorax with narrow scales as well as hairs ; last. two white rings of
female palpi rather broad .. es oh super pictus, Grassi.

8. Thorax with hairs only ; male aedoeagus Sh leaflets
(turkhudt, Theo.), bees Theo.
Sides of mesonotum with distinct narrow scales : $
9. Male aedoeagus with leaflets .. — = neh var. persicus, n.
Male aedoeagus without leaflets Ke Ne as .. multicolor, Camb.
Larvae.

1. Outer clypeal hair branched... ts ay > ah ae 5 Sane
Outer clypeal hair simple se Bh ei: 3 MTSE ie oS

2. Palmate tufts on abdominal segments i-—vil; outer clypeal hair thickly
branched ae . pharoensis, Theo.

Palmate tufts on abdominal segments lli—Vvii ; outer clypeal hair less
branched a6 : oe m pulcherrimus, Theo.

. A rudimentary palmate tut on ae st abdominal segment

costalis, Theo. ; rhodesiensis, Theo.
This tuft absent .. a% tre 55 ae ia a A pees:

. Posterior clypeal hair short ; innermost shoulder-hair much branched

stephenst, Liston.
Posterior clypeal hair long

. Innermost shoulder-hair much br od miter ke superpictus, Grassi.

Innermost shoulder-hair slightlv branched towards tip |
multicolor, Camb. ; 2? hisbaniola, Theo. ; ? turkhudi, Theo.

Good distinctions between some of the above larvae have yet to be discovered ;
possibly some may be found in the markings of the head. The larvae of A. rhodesiensis,
A. hispaniola and A. turkhudi have not been fully described, and the British Muséum
does not possess sufficient material from which to supplement the descriptions.

MOSQUITOS OF THE PALAEARCTIC REGION. 277

10. Anopheles (Myzomyia) pulcherrimus, Theo. (fig. 2, g).

Anopheles pulcherrimus, Theobald, Proc. R. Soc. Ixix, p. 369 (1902).

A very well-marked species, easily distinguished from all others in the Palaearctic
fauna, except A. pharoensis, by having the abdomen densely covered with broad
flat scales, rather loosely applied and forming distinct lateral tufts. The front
femora are slightly swollen near the base, and in this point, as well as in the shaggily
scaled female palpi, A. pulcherrimus and A. pharoensis seem to show some approach
to A. hyrcanus; but the type of wing-markings and the structure of the male
hypopygium are typical of the subgenus Myzomvyra.

The early stages have been described by Vassiliev.

Distribution.—Semi-arid regions of south-western Asia, from Turkestan and the
Punjab to Lower Mesopotamia.

11. Anopheles (Myzomyia) pharoensis, [heo.

Anopheles pharoensis, Theobald, Mon. Cul. i, p. 169 (1901).
Anopheles maculicosta, Becker, Mitt. Zool. Mus. Berlin, ii, 2 '6o. (1903),

Very similar to A. pulcherrimus, but is somewhat larger and has a broad dark
ring on the fourth hind tarsal joint ; usually also there is a pair of conspicuous black
spots about the middle of the mesonotum, which are not seen in A. pulcherrimus.
In both species the wing-membrane is deeply stained in the areas occupied by the
main patches of dark scales. A similar condition is seen in some American. species,
and to some extent discounts Christophers’ theory that all the pale markings of
Anopheles wings arose by bleaching from a primitive, uniformly dark condition.

Distribution —Essentially an Ethiopian species, being widely spread, though
not common, in tropical Africa, and occurring in Madagascar. Common in Lower
Egypt, and occurring rarely in Palestine (Barraud).

12. Anopheles (Myzomyia) stephensi, Liston (fig. 2, h).
Anopheles stephensi, Liston, Ind. Med. Gaz. xxxvi (1901) ; Christophers, Ind.
J. Med. Res. iii, p. 481 (1916).

The femora and tibiae are conspicuously spotted with white, and the abdomen
is almost covered with scales; the scales are narrow, and less dense than those of
A. pharoensis and A. pulcherrimus, and never form lateral tufts. None of the
allied Oriental or Ethiopian species (A. maculatus, A. willmori, A. maculipalpis,
A. theobaldi, etc.) appear to have occurred within the area under discussion, and
A. stephensi need not therefore be compared with them for our present purpose.

Distribution.—An Indian species occurring in a localised area round Basra, at
the head of the Persian Gulf (Barraud and Christophers).

13. Anopheles (Myzomyia) costalis, Theobald (fig. 2, i).

Anopheles costalis, Theobald, Mon. Cul. i, p. 157 (1901).
? Anopheles costalis, Loew, Berlin. ent. Zeitschr., x, p. 55 (1886).
Anopheles arabiensis, Patton, J. Bombay Nat. Hist. Soc. xvi, p. 625 (1905).

The spots on the femora and tibiae vary in number and intensity, but are always
present, and together with the absence of abdominal scales render the species an
easy one to identify when Palaearctic forms alone are considered. The female
palpi have three whitish rings, the terminal one broad and including the whole of
the last joint.

I am informed by Dr. Enderlein that Loew’s type of A. costalis is not now in his
collection in the Berlin Zoological Museum.

21S F. W. EDWARDS.

Distribution.—Essentially an Ethiopian species, througnout which region it is
abundant. I only include it among the Palaearctic species on the strength of a
single female in the Paris Museum captured in Algeria (locality not noted) by
M. E. Roubaud. It does not appear to have been recorded from Egypt, though
there seems no reason why it should not spread there from the south.

14. Anopheles (Myzomyia) rhodesiensis, Theo. (fig. 2, j).

Anopheles rhodesiensis, Theobald, Mon. Cul. i, p. 184 (1901).

Anopheles d’thali, Patton, J. Bombay Nat. Hist. Soc. xvi, p. 627 (1905).

This species could not be confused with any other in the Palaearctic fauna,
although the darkest varieties of the African A. funestus approach it rather closely.
A detailed description nas been given by Christophers (1915), who records it from
Arabia, Baluchistan and Quetta. Its occurrence in Baluchistan is the only reason
for including it in this paper. It occurs also in widely separated parts of tropical
Africa.

15. Anopheles (Myzomyia) superpictus, Grassi (fig. 3).

Anopheles superpictus, Grassi, Reale Accad. Lincei., p. 78 (1900).

Pyretophorus palestinensis, Theobald, Mon. Cul. iu, p. 71 (1903).

Pyretophorus nurset, Theobald, Mon. Cul. iv, p. 66 (1907).

Pyretophorus cardamitisi, Newstead & Carter, Ann. Trop. Med. iv, p. 379 (1910).

Anopheles superpictus var. macedoniensis, Cot & Hovasse, Bull. Soc. Path. Exot.

<p. Soo tS 7):

Distinguished from the nearly allied A. multicolor by the broad white tips to the
female palpi. In both species the scutum bears distinct scales, at least at the sides.
The wing markings of A. superpictus are very variable, the pale costal spots being
sometimes much reduced. According to Ficalbi the tarsi are faintly ringed with

Fig. 3. Larva of Anopheles superpictus, Grassi: a, palmate tuft; 56, mentum.

pale at the joints, when viewed through a lens. This is certainly true of a few of the
specimens I have examined, but the majority have the tarsi entirely dark. I think,
however, that there is no doubt that A. superpictus and A. palestinensis are the
same species.

———

MOSQUITOS OF THE PALAEARCTIC REGION. 279

The larva has not been fully described. It may be diagnosed as follows :—
Antennae with a minute single hair before the middle. Frontal hairs all simple,
and all about equal in length, the inner anterior pair widely separated, the posterior
pair almost immediately behind the inner anterior. Head markings fairly constant,
arranged as in the figure, the spots small. Innermost shoulder hair plumose
from base to tip, the hair next to it also plumose, but longer and with fewer branches.
No rudimentary palmate hairs on thorax or on first abdominal segment ; a pair of
small ones on the second segment, and fully developed pairs on segments 3-7 ; each
fully developed hair has about 16 leaflets, which are lanceolate, with two or three
jags beyond the middle, forming a rather indefinite shoulder ; terminal portion
long, but not nearly as long as the basal. No difference could be detected between
Macedonian and Indian specimens.

The species is said to be associated with mountainous districts, though by no
means confined to high altitudes, and to be largely a stream-breeder.

Distribution.—Eastern Mediterranean region, from Italy to Macedonia and
Palestine ; thence through Upper Mesopotamia and Persia to the Punjab.
Also recorded from North Africa, though I have seen no specimens from there.
Transcaspia (Firudza, C. Ahnger).

16. Anopheles (Myzomyia) sergenti, [heo.

Pyretophorus sergenti, Theobald, Mon. Cul. iv, p. 68 (1907).

This species very much resembles A. superpictus, but 1s rather smaller, and
differs as indicated in the key. I have previously considered it identical with the
Indian A. culicifacies, but whereas in A. sergenti there are four or five pale spots
in the wing-fringe, in A. culicifacies there is never more than one. A. sergentt
might equally well be regarded as a variety of the African A. funestus, which has
several pale fringe-spots, but is much darker in general coloration, and has the third
vein much less extensively pale. Detailed comparisons of these three species in all
their stages are necessary before final conclusions can be reached as to their exact
relationships.

Distribution.—Algeria (Sergent) ; Tunis (Tamerza, Dr. M. Langeron) ; Palestine
(Annandale, Barraud).

17. Anopheles (Myzomyia) hispaniola, Theobald (fig. 2, m).

Myzomyia hispaniola, Theobald, Mon. Cul. ii, p. 49 (1903).
Pyretophorus myzomyfacies, Theobald, Mon. Cul. iv, p. 69 ( 1907).

Apparently only differs (in the adult) from A. multicolor in having leaflets present
on the male aedoeagus, and hairs instead of narrow scales on the scutum ; according
to Sergent & Foley the eggs of the two species are remarkably distinct. The wing-
markings of both species (especially A. multicolor) are so variable that no constant
difference can be found between them. A. /ispaniola is quite indistinguishable,
so far as I can see, from the Indian A. turkhudi in the adult state. The egg of
A. hispaniolaas figured by Sergent is, however, so different from that of A. turkhudi
as figured by Christophers that, unless a mistake has been made by either author,
it is impossible to regard the two as conspecific, especially as they are widely discon-
tinuous in their distribution. The larvae also differ, according to the published
descriptions, in the number of pairs of palmate hairs present ; but this requires
confirmation, since the figure given by James and Liston is inconsistent with their
description.

This species is apparently much less fond of desert conditions and saline water
than A. multicolor.

280 F. W. EDWARDS.

Distribution.—S. Spain (Macdonald); Algeria (Sergent); S. Tunis (Tamerza,
Dr. M. Langeron). Does not appear to occur in Egypt or Palestine.

References.—Sergent (Ed. & Et.), Ann. Inst. Pasteur, xix, 1905, p. 144 ; xx, 1908,
p. 393.

18. Anopheles (Myzomyia) turkhudi var. persicus, nov.

The type male of A. turkhudi has no scales on the scutum, and has distinct
leaflets on the aedoeagus. The same is true of all specimens I have seen from the
Punjab, but a number sent me by Christophers from East Persia show distinct scales
on the scutum, and are in fact indistinguishable externally from A. multicolor.
They can hardly be that species, however, since the male aedoeagus bears distinct
leaflets, as in A. turkhudi and A. hispaniola. Provisionally, therefore, I regard them
as representing a distinct variety of A. turkhudi, which inhabits adjacent areas in
the Punjab. The real relationships of all these forms can only be determined by a
close study of their habits, early stages and distribution, and the early stages of this
Persian form are not yet recorded.

19. Anopheles (Myzomyia) multicolor, Camb. (fig. 2, k).
Anopheles multicolor, Camboulin, C. R. Acad. Sci. exxxv, p. 704 (1902).
Pyretophorus chaudoyei, Theobald, Mon. Cul. iii, p. 68 (1903).
Pyretophorus cleopatrae, Willcocks (nom. nud.).
Anopheles impunctus, Doénitz, Zeitschr. f. Hygiene, xli, p. 67 (1902).

The adult does not differ appreciably from A. hispaniola and A. turkhudi except
in having scales on the mesonotum (these scales, however, varying in number, width,
and distinctness), and in the entire lack of leaflets on the male aedoeagus. 4A.
superpictus has a similar thoracic ornamentation, but can easily be distinguished
by its white-tipped palpi. Egyptian specimens commonly have the wings more
extensively dark than those from Algeria, but, on the other hand, it is almost certain
that Donitz’s A. impunctus, described from an abnormally pale specimen, is only
a variety of this species.

The larva has been fully described by Foley (1912) and Langeron (1918).
According to Langeron’s figures the innermost shoulder-hair is only trifid at the
tip, but in a number of specimens he has sent me from Tozeur, as well as in some
skins sent by Capt. Barraud from Palestine, it is branched almost from the base
and somewhat plumose. The head of the larva is extremely dark, the usual markings
being all fused in most specimens into a large black patch which occupies the greater
part of the head, leaving the front of the clypeus yellowish. The lower surface of
the head, as well as the upper, is mainly blackish.

The egg as described and figured by Foley (reproduced in fig. 2, k) is very dis-
tinctive, and quite unlike that of A. hispaniola or A. turkhudi. The species is said
to breed in highly saline desert pools, or in brackish water near the sea.

Distribution.—Desert regions of North Africa from Southern Algeria to Egypt ;
Palestine ; Teneriffe. Not certainly known from further east or north.

Tribe CULICINI.

I propose to revert to the old classification of mosquitos, and recognise only two
tribes, the ANOPHELINI and CuLIciNI, including in the latter the MEGARHININI and
SABETHINI. I am now convinced that Howard, Dyar and Knab were right in
including Megarhinus in the Curicrnt and placing it somewhere near Psorophora ;
the modifications of the adults, though striking, are not fundamentally important.
On the other hand, I am equally convinced that the tribe SABETHINI cannot be
maintained. Every character on which it has been attempted to define it breaks

MOSQUITOS OF THE PALAEARCTIC REGION. 281

down at some point or other. The character finally adopted by Howard, Dyar and
Knab, that of the head bristles, is not indicated in Evetmopodites, which is unquestion-
ably a Sabethine genus, while it is shown in some Culicine genera, notably Megarhinus.
There is, no doubt,a group of genera (or more probably two or three independent
groups) which are more closely related to one another than to other Culicines, but
in adult structure there is certainly no hard and fast line to be drawn. In regard
to the absence of the anal brush of the larva, which is the one common and out-
standing feature of all SABETHINI, Colonel Alcock has suggested to me (I consider
with great plausibility) that this may be an adaptive character, and therefore not
necessarily indicative of relationship. As he points out, larvae living in small
confined spaces, as do almost all of the SABETHINI, would not require the anal brush
for swimming ; and as it is absent in the first stage, there seems no reason whatever
why its development should be proceeded with in the later stages. In confirmation
of this view, it may be remembered that some of those species of Aédes (Stegomyta)
which live in small collections of water have the brush much less developed than in
the free-living forms.

The ancestral form probably had pulvilli and spiracular bristles, narrow scales,
and a non-carnivorous larva ; partaking thus of some of the characters of Theobaldia
and Lutzia ; no such form is known, however.

Table of Genera of Palaearctic Culicine Mosquitos.

Adults.
1. Vein A, (6th) ending below or a little before the level of the base of R,
(2nd) ; no microtrichia on wing-membrane us ie Uranotaenia.
Vein A, ending well beyond the level of the base of R,; wing mem-
brane with microtrichia .. a ies ee ue Me 2 aia

2. Pulvilli absent 3
Pulvilli present oe x ie af - * cd
3. Spiracular bristles present 5 = i Bs nbs :s nr
Spiracular bristles absent ‘ ede eS)

4. Proboscis slender, flexible, as long as tie sitole ede ; one pro-epimeral
bristle (or none); no sternopleural bristles. is .. Rachionotomyia.
Proboscis otherwise ; sternopleural bristles present ab oe af 55 feo

5. Proboscis rigid, hooked; no pro-epimeral bristles; cell R, extremely

short ; a V-shaped ‘thickening of the wing-membrane between

veins Cu, and Cu, Gorksiofrothye 2 ‘ : Megarhinus.
Proboscis flexible, straight in repose ; several pro- epimeral bristles ; cell

R, long; no V-shaped thickening of wing-membrane. . . Theobaldia.

6. Two pro-epimeral bristles ss be ay a Cie Sion

Several (about 5) pro-epimeral br istles. oF an ae ae a ame,

7. No post-spiracular bristles ; female claws simple
Taentorhynchus (subgenus Coquillettidia).
At least a few small post-spiracular bristles present ; female claws nearly
always toothed

8. Proboscis not very stout, eiaient or narged ee in repose. A édes.
Proboscis stout, Tipu (from dry specimens) curved downwards in

GEPOSe.. . : ‘ ais i a au .. Armigeres.

9. Lower mesepimer al thstles numerous. ge a2 tis Lutza.

Normally one lower mesepimeral bristle ; often none, but very rarely
two or three .. es ee ai ae 8 ae SF Culex.

iS)
(o2)
bo

F. W. EDWARDS.

Larvae.

1. Anal brush and barred area absent ; thorax with a strong spine at each
posterior corner ns : .. Rachionotomyia.
Anal brush and barred area pr esent (except i in ‘first stage) 3 oe pe eee
2. Siphon with one pair of ventral tufts .. oe es a 5g en:
Siphon with several ventral tufts i: oF ae ae ss ee
3. Siphonal tuft at or near base : ss ee ate 4
Siphonal tuft near middle, or bey ond Se ; 5 )

4. Mouth-parts modified for predacity ; a chitinous eee on oat side of
eighth segment ; body colour dark reddish 43 Megarhinus.

Mouth-parts not modified for predacity ; a patch of scales on each side
of eighth segment ; body colour not reddish oe tk Theobaldia.

5. Pecten teeth rounded and fringed apically ; a chitinous plate on each side
of eighth segment (in fourth stage only) in addition to the comb

Uranotaema.

Pecten teeth (when present) sharply pointed, with denticles on one
side; eighth segment with comb or patch of scales... 3 Rend ae)

6. Valves of siphon and_ tracheal ae highly modified for subaquatic
respiration... : .. Laentorhynchus.
Valves of siphon and tracheal sy stem not specially modified ae way, a

7. Extremely long single hairs on thorax and abdomen, besides long hair
tufts ; dorsal chitinous plates on segments 6-8 in the fourth stage ;

colour reddish . : .. Orthopodomyia.
Single hairs not longer than the moderate tufts ; no dorsal chitinous
plates on segments 6-8 ; colour not reddish Me fs Pe ee ue:
8. Pecten present .. oe Kis es a =e ws bd Aédes.
Pectenvabsent: ~~. : ar Armigeres.
9. Mouth-parts modified ue pr sceaiete ; anal segment iene and aod Lutzia.
Mouth-parts not modified ; anal segment shorter and less pointed. . Culex.

Genus Uranotaenia, Arrib.

Most writers have distinguished this genus mainly by the short upper fork-cell
(cell R,), but while this is a sufficient distinction in the majority of cases, there are a
few species (e.g., U. unguiculata) in which the shortening of this cell is not very
noticeable, and, on the other hand, some species of the Aédes group have the cell
so short that they have been mistaken for species of Uranotaenia. A more absolutely
diagnostic character, though requiring a high magnification for its detection, is the
absence in all known species of the genus of microtrichia on the wing-membrane.
This distinguishes Uvanotaenia sharply from all other CuLicrpar. The short anal
vein, ending below or before the base of the radial sector, is shown also by the tropical
genera Hodgesia and Harpagomyia. In all known species except U. wnguiculata
the front claws of the male are small and equal in length. The pleural bristles are
very much reduced in number, there being only one or two in each of the pronotal,
pro-epimeral, spiracular and pre-alar series.

The larva is not very sharply distinguished from that of Aédes, except in the
fourth stage, when the characteristic lateral chitinous plates appear on the eighth
abdominal segment. The spine-like frontal hairs found in many species are repre-
sented by normal simple hairs in others (¢.g., U. unguiculata). The form of the
pecten-teeth is characteristic. The resting position in the water is almost horizontal.

The genus is tropicopolitan, but two species occur within the limits of the Palaearctic
region.

MOSQUITOS OF THE PALAEARCTIC REGION. 283

1. U. unguiculata, Edw. .
Uranotaema unguiculata, Edwards, J. Proc. Asiatic. (Soc.) Bengal, ix, p. 51 (1913).

A line of pale blue flat scales passes round the margin of the mesonotum from
wing-base to neck. The front claws of the male are unequal, the larger one with a
small tooth.

The larva has been described by Joyeux.

Distribution.—Eastern Mediterranean region. Originally described from North
Palestine, it has since been found in Jerusalem (Goldberg), Egypt (Gough), Macedonia
(Waterston, Joyeux), and Italy (Hargreaves), but is always rare.

2. U. bimaculata, Leicester.

Uranotaenia bimaculata, Leicester, Cul. of Malaya, p. 226 (1908).

There is no line of flat scales on the margin of the mesonotum, but just in front
of each wing-base is a large oval velvet-black spot on the integument. The front
claws of the male are normal for the genus, small, equal and simple.

The larva is not yet described.

Distribution—Japan (Tokio, S. Yamada); Malay Peninsula (Levcester). There
is also a representative species (U. mashonaensis, Theo., = U. bimaculata, Theo.)
in tropical Africa, which scarcely differs from the Oriental form. Should the two
be united, the species must be known as U. mashonaensis, Theo.

Genus Rachionotomyia (Theo.) Edw.

This genus has never been fully described. I would define it as follows :—

Proboscis slender throughout ; longer than the long front femora, and as long
as the whole body. Mouth-parts normal; maxillary teeth very small. Palpi
short in both sexes, not more than one-sixth as long as the proboscis. Female antennae
slender, the verticils about four times as long as the joints; male antennae with
shorter joints (except the last two) and longer verticils. Eyes practically touching
above the antennae, the supra-antennal portion broad. A pair of strong bristles,
placed close together, projecting forwards from immediately above the line of contact
of the eyes ; far away from these bristles are two other smaller ones on each side of
the head. Prothoracic lobes well separated. Mesonotum with or without well-
developed dorso-central bristles. One pro-epimeral bristle (even this is absent in
some species). Several (3-6) spiracular bristles. No postspiracular bristles; no
row of bristles on the upper part of the sternopleura ; no lower meso-epimeral bristles.
Sub-alar knob small, with few bristles. Postnotum bare, with a pair of slight furrows
dividing it into three portions. Last segment of female abdomen blunt, very bristly.
Male hypopygium: Lobes of ninth tergite elongate, with long apical bristles ; side-
pieces rather short, with conical, bristly claspette lobes; clasper long, terminal,
with small thick terminal claw; tenth sternites split apically into several short,
tergally projecting teeth, in a longitudinal row; aedoeagus very small, a simple,
incomplete tube, with small parameres and sometimes apparently with some internal
spines. Hind tibiae somewhat shorter than the others. Female claws simple ;
front claws of male unequal. Pulvilli absent. Wing-membrane with distinct
microtrichia. Rg somewhat longer than R,, 3; fork-cells moderately long; anal
vein reaching well beyond the level of the base of Rx.

Larva (description based on a comparison of three Oriental species) : Antennae
short, smooth, apparently without hair on shaft. Clypeal hairs very small, simple.
Posterior corners of metathorax with a long strong spine situated on a chitinous
plate, without any accompanying long hairs; a similar but smaller spine towards
each side of the mesothorax. First seven abdominal segments with seven pairs of
strong stellate tufts (three dorsal, two lateral, two ventral). Thorax and first six

284 F. W. EDWARDS.

abdominal segments with long, lateral plumose hairs. Comb of eighth segment a
row of stout spines, sometimes attached to a plate. Anal plate bears a comb of a
few short spines. Siphon moderate, with sparse latero-ventral pecten and numerous
hair-tufts or single hairs. Pupal paddles small, somewhat pointed, without fringe
or terminal hair. Habitat: pitcher plants, bamboos, etc.

The genus is characteristic of the Oriental and Australasian regions, one species
occurring in Japan.

Fig. 4. Male hypopygium of Rachionotomyia bambusa, Yam.: a, ventral view, x75; }, lateral
view, x75; c, aedoeagus, ventral view, x225.

Rachionotomyia bambusa, Yamada (fig. 4).
Rachionotomyia bambusa, Yamada, Dobutz. Z. Tokio, xxix, pp. 61-72 (1917).

This belongs to the rather numerous group of species with metallic silvery markings
on the pleura and abdomen, and silvery spots on the femora. The chief diagnostic
characters are as follows :—Head with a broad blue band in front. Prothoracic
lobes with black scales. Pro-epimera with a few narrow black scales and one bristle.
Mesonotum almost entirely dark brown, with long, straight, very narrow greenish-
black scales. Scutellar scales broad, greenish black. Integument of pleurae
mainly dark brown. Larger claw of front legs of male thick, with a pointed swelling,
scarcely a tooth, beyond the middle. Lobes of ninth tergite of male each with about
nine long, rather stout bristles.

I have not seen Yamada’s description of the larva, and have had no specimens
for examination.

Distribution.—Japan (Tokio, S. Yamada ; Kofou, L. Drouard de Lezey).

Genus Megarhinus, k.D.

I now feel strongly inclined to accept the view of Howard, Dyar and Knab that
Toxorhynchites should not be separated from Megarhinus, except as a rather
weakly-marked subgenus. There are no important structural differences whatever,
Toxorhynchites differing only in the shorter female palpi.

As Banks has pointed out, the labium in this genus is a rigid organ, and the
structure of its apical part is apparently such as to prevent the possibility of the
mouth-parts being used for sucking blood.

The pleural bristles in this genus are remarkable; there appear to be no true
pro-epimeral bristles, but, on the other hand, a row of strong spiracular bristles is
developed ; there are no post-spiracular bristles, and the only bristles on the pro-
thoracic lobes are a few which point forwards or downwards. There are only two
pairs of ocular bristles instead of the usual row. The reduction of the mesonotal
bristles has been remarked on by previous writers.

MOSQUITOS OF THE PALAEARCTIC REGION. 285

The presence of spiracular bristles, the close similarity in the male hypopygium,
and the basally situated siphon-tuft of the larva, are points which seem to indicate
some connection, distant no doubt, with Theobaldia, a connection which would
not be suspected on a superficial examination.

The species of this genus often have very specialised habits, and these have
a restricted distribution. Two species have already been recorded from the
Palaearctic region ; what is probably a third exists in south-east Europe.

1. Megarhinus christophi, Portch.
Megarhinus christophi, Portchinsky, Horae Soc. Ent. Ross, xviii, p. 122 (1884).

I do not know this species. According to Portchinsky’s description it differs
from M. towadensis in having silvery transverse bands on each of the first six abdominal
segments and yellow lateral hairs on the sixth segment.

Distribution.—Amur (Portchinsky).

2. Megarhinus towadensis, Mats.

Megarhina towadensis, Matsumura, Thousand Insects of Japan, Add, ii, p. 445
(1916).

This is very much like the common Oriental M. regius (Tennent), but there are
differences, evidently of specific value. The only specimen I have seen is one sent
for determination from the Paris Museum by M. Séguy. This agrees with Mat-
sumura’s description, except that the narrow white ring on the first joint of the
middle tarsus is close to but not at the base (the front legs are missing). The thorax
is much rubbed, but the remaining mesonotal scales are metallic emerald green,
quite unlike the dull scales of M. vegius. The abdominal tuft is large and conspicuous,
the long hair on the sixth and seventh segments black, on the eighth segment orange.
The venter is mainly shining blue, but there are lateral yellow patches on
sternites two, three and six, and sublateral, elongate yellow triangles on sternites
four and five.

Distribution.—Japan (Towada, Matsumura ; Kofou, L. Drouart de Lezey, 1906).

3. Megarhinus sp.

It is necessary to mention here a Megarhinus larva which was obtained near
Karasouli, Macedonia, by Capt. Waterston.

The specimen was taken with a number of other larvae of different insects which
were thought to be preying on mosquito larvae. It was not identified until after
the collector’s return to England, and no attempt was made at the time to rear it
or to obtain others. It differs only in minute characters from the larvae of the
African M. brevipalpis, the Oriental M. regius, and the North American M. rutilus,
and it is therefore impossible to say whether it belongs to one of these species, or to
some other, perhaps undescribed, form.

Genus Theobaldia, Neveu-Lemaire.

The two characters on which most writers have based their conception of this
genus—the spotted wings and the position of the cross-veins in a straight line or
nearly so—are both worthless for purposes of definition. In T. longiareolata and
in the subgenus Culicella the cross-veins are well separated ; while, on the other hand,
in Lutzia vorax and occasionally in some other species (¢.g., Culex tipuliformis,
Aédes rusticus) they are practically in a straight line. The female palpal character
on which Neveu-Lemaire founded the genus is of even less value. In one point,
however, the adults do seem to be sharply distinguished from most other genera of
the tribe Cuticrni. This is in the possession, immediately in front of the prothoracic

(4183) x

286 F. W. EDWARDS.

spiracle, of a row of yellow bristles.* These bristles are quite apart from the usual
pro-epimeral bristles, which are present in Theobaldia but somewhat reduced in length,
their place being largely taken by the spiracular bristles, which are placed in a close-
set row on the posterior side of the posterior ridge of the pro-epimeron. As in
Orthopodomyia, there are no post-spiracular bristles.

The African species, Leptosomatomyia fraseri, shows the same pleural and hypo-
pygial characters as Theobaldia, and should probably be included here. Apart from
this, the genus is practically confined to the temperate regions of the northern
hemisphere, three species occurring in the Punjab.

The larvae are sharply distinguished from others in the Palaearctic fauna
(except Megarhinus) by the basally situated hair-tuft on the siphon.

Three subgenera (Theobaldia, Culicella and Allotheobaldia) are sharply defined
by larval structure, but the distinction in the adults is not so clear, and it will therefore
be best to tabulate all the species together, while admitting the subgenera as valid.
I agree with Brolemann that Culicella is not generically separable from Theobaldia.

Adults.

1. Thorax with rather sharply defined white lines; femora and tibiae
striped and spotted ; costa largely pale-scaled ; male palpi shorter than

the proboscis (subgenus Allotheobaldia) .. ss longiareolata, Macq.
Thorax without sharply defined white lines; femora and tibiae not

striped ; costa all dark ; male palpi longer than the proboscis sixe te ee

2. Cross-veins in a straight line, or nearly (subgenus Theobaldia) .. sues bl
Cross-veins well separated ; tarsi with narrow pale rings embracing both

ends of joints ; wings not distinctly spotted (subgenus Cudicella) ee es

3. Tarsi entirely dark ; wings not distinctly spotted .. glaphyroptera, Schin.
Tarsi broadly ringed with white at the bases of the joints ; wings more
distinctly spotted (at least in the female) by accumulation of scales at

the bases of the fork-cells and on the cross-veins a a a: 4
4. Femora without pale pre-apical ring; hind metatarsi without white ring
in the middle.. : ; ik alaskaensis, Ludlow.

Femora with pale pre-apical ring ; hind metatarsi with white ring in middle 5

5. Wing-spots distinct ; abdomen conspicuously banded with black and
white .. ane ae a a A me .. annulata, Schrank.

Wing-spots indistinct, especially in the male; abdominal scales nearly
all ochreous... a es = o3 es .. subochrea, Edw.

6. First joint of front tarsi of male longer than the remaining joints
together ; proboscis of female almost entirely black .. —morsitans, Theo.

First joint of front tarsi of male not longer than the remaining joints

together ; proboscis of female with many pale scales at the sides and
beneath 5 ee ts sts ae ae fumipennis, Steph.

Male Hypopygia.

1. Ninth tergite with a pair of long processes ; aedoeagus remarkably large
and complicated ae A a : longiareolata, Macq.
Ninth tergite without distinct processes; aedoeagus normal .. Seas 4

. Clasper somewhat swollen at the tip ; side-piece with a subapical projection
bearing a number of long, flattened appendages .. — glaphyroptera, Schin.
Clasper not swollen at the tip; side-piece otherwise a bye Nh
3. Side-piece with a more or less distinct, hairy subapical knob sis a
Side-piece without subapical knob or agglomeration of hairs 5

* This character is also possessed by Megarhinus, Psorophorva and some Sabethine genera
(e.g., Rachionotomyia), but none of these could well be confused with Theobaldia.

MOSQUITOS OF THE PALAEARCTIC REGION. 287

4. Tip of eighth sternite with a close-set row of short spines alaskaensis, Ludlow.
These spines absent Ag .. annulata, Schrank. ; subochrea, Edw.
5. Side-pieces barely three oe as long as their basal width ; ap of eighth
sternite without row of spines : .. morsitans, Theo.
Side-pieces four times as long as their basal width ; tip of eighth sternite
with row of short spines oe Sf: as om fumipennis, Steph.

Larvae.
1. Antennae short, with small hair-tuft or a single hair; siphon short, at
most three times as long as broad . a ee he ne
Antennae long, with large, many- -branched tuft ; siphon long, at least five
times as long as broad (subgenus Culicella) by :
2. Siphon with 6-10 stout, widely-spaced spines (subgenus i ae
longiareolata, Macq.
' Siphon with a well-marked pecten, most of the teeth of which have their

tips drawn out into long hairs (subgenus Theobaldia)
annulata, Schrank ; subochrea, Edw.
3. Pecten straighter ; siphon without accessory spines .. morsitans, Theo.
Pecten oblique ; siphon also with accessory spines .. . .fumipennis, Steph.

The larvae of T. glaphyroptera and T. alaskaensis are unknown.

Subgenus Allotheobaldia, Brolemann.
1. Theobaldia (Allotheobaldia) longiareolata (Macquart).

Culex longiareolatus, Macquart, Dipt. Exot. i, 1, p. 34 (1838).
Culex spathipalpis, Rondani, Bull. Soc. Ent. Ital. iv, p. 31 (1872).
Culex serratipes, Becker, Mitt. Zool. Mus. Berlin, iv, p. 78 (1908).
Culex annulatus var. marocanus, d’Anfreville, Bull. Soc. Path. Exot. ix, p. 140
(1916).
A very distinct species, which could not be confused with any other, either in
the adult or larval state. The larva has been fully described by Langeron.

Distribution Throughout the Mediterranean region and in most of the Atlantic
islands ; spreading southwards by East Africa as far as the Cape Province, and
eastwards through Transcaspia, Mesopotamia and Persia to the Punjab. In France
it has been taken as far north as Rambouillet (Villeneuve), but it appears to be other-
wise unknown in central and northern Europe. Some new records are: Transcaspia
(Askhabad, Firudza, Vrefskaja, A/mnger) ; Seistan (Annandale).

Subgenus Theobaldia, Neveu-Lemaire.
2. Theobaldia (Theobaldia) glaphyroptera (Schiner).

Culex glaphyropterus, Schiner, Fauna Austriaca, ii, p. 628 (1864).
Theobaldia bergrothi, Edwards, Entom. Tidskr. p. 50 (1921).

This was described by Schiner and Ficalbi as having spotted wings, and assuming
that these authors’ statements indicated a definite spotting such as that of T. annulata,
I was led to regard the specimens sent me by Drs. Bergroth and Frey as a distinct
new species. Later, however, I received a series of specimens from the Vienna Museum,
including some of Schiner’s original series, and these showed that the spotting of the
wings is really hardly distinguishable, except for a darkened area of the membrane
towards the costa in the middle. The Finnish and Swedish female specimens for
which I proposed the name bergrothi are almost certainly only T. glaphyroptera.
On the other hand, Theobald’s Pseudotheobaldia niveitaeniata, which I supposed in
1913 to be synonymous with T. glaphyroptera, is a perfectly distinct species.

(4183) XZ

288 F. W. EDWARDS.

T. glaphyroptera is distinct in coloration from all other European species, but is
very similar to the North American T. impatiens, Walk., from which it differs by the
very distinctive male hypopygium. The male palpi are much more slender than
those of any other European species, and seem to indicate an approach to Culex,
although the terminal joints are not upturned. The omamentation of the thorax
is variable, some specimens having a distinct pattern of ochreous or whitish lines
(one straight median, and a pair of curved lateral) on a dark ground.

The larva is unknown.

Distribution.—Recorded by Schiner from Austria, and by Eckstein from the
Strasburg district. The specimens I have seen are from Austria (Rekawinkel, Pokorny ;
Hammern, Mik; also several from Schiner’s original series, collected by Egger) ;
Moravia (Frain, Handlirsch); Hungary (Bartfa, Kertész); Sweden (Stockholm ;
Dalecarlia, Norrbotten, Boheman ; Ostergétland, Haglund); Germany(?) (Coll.
Loew, Berlin Museum); Finland (Kivikoski, Saima-Canal, Adelung; Kuustd,
Lundstrom ; WKittila, Krogerus ; Petrosaw, Giinther ; Sarasniemi, Wuorentaus).

3. Theobaldia (Theobaldia) alaskaensis, Ludlow.

Theobaldia alaskaensis, Ludlow, Can. Ent. xxxviii, p. 326 (1906).
Culiseta siberiensis, Ludlow, Insec. Inscit. vii, p. 151 (7th Jan. 1920).
Theobaldia arctica, Edwards, Bull. Ent. Res. x, p. 136 (end Jan. 1920).

In describing T. arctica I compared it with T. alaskaensis, and suggested that
the two might be the same, though according to the published figure of 7. alaskaensis
there appeared to be certain differences in the male hypopygium. Recently Dr.
H. G. Dyar sent me a male from Alaska which shows that the supposed differences
do not exist. The species is easily distinguished by leg markings from the other
European forms, but is represented by allied species, differing in hypopygial structure,
in Mexico and the Punjab.

Distribution—I have examined specimens from Archangel; Scotland; N.
England; Sweden (Norrbotten, Boheman; Jemtland, Schénherr ; Stockholm,
Boheman) ; Lower Austria; Upper Silesia (Astron, Brauer); Siberia (Irkutsk,
Ahnger ; Inserovo, 62° 5’, Trybom). Ludlow and Dyar record it from Eastern
Siberia, Alaska and Alberta.

4. Theobaldia (Theobaldia) annulata (Schrank) (fig. 1A).

Culex annulatus, Schrank, Beitr. Z. Naturg. p. 97 (1776).

? Culex variegatus, Schrank, Enum. Ins. Austr. p. 482 (1781).
? Culex annulatus, de Fourcroy, Ent. Paris, p. 516 (1785).
Culex annulatus, Fabricius, Mantissa Ins. ii, p. 363 (1787).

? Culex nicaensis, Leach, Zool. Journ. ii, p. 292 (1825),

Culex affinis, Stephens, Zool. Journ. iv, (1825).

A common and easily recognised domestic species, normally exhibiting very
little variation except in size. North European specimens are on the average larger
than those from the south.

The name Culex annulatus was apparently proposed independently by Schrank,
de Fourcroy and Fabricius, but the same species was designated in each case. De
Fourcroy’s diagnosis is totally inadequate, but he gives the habitat as Parisian
gardens, so that he more probably had this species than an Aédes.

C. variegatus, Schrank, is generally quoted as a synonym of T. annulata, but the
inadequate diagnosis would perhaps serve better for Aédes lutescens. Walker
mentions specimens of 7. annulata in the British Museum ‘‘ from Dr. Leach’s
collection,’ and it 1s possible that these specimens (which are no longer in existence)
may have been the originals of C. nicaensts.

The larva has been described by Langeron.

MOSQUITOS OF THE PALAEARCTIC REGION. 289

Distribution —Throughout Europe, but probably commoner in the north than
in the south, where it seems to be largely replaced by T. longtareolata ; extending
into Palestine (Jerusalem, Dr. Goldberg) and North Africa (Biskra, Algeria, Eversmann ;
etc.). Has not yet been found further east, but is represented in the United States
by an allied species. A female in the Berlin Museum is labelled “ Ural, Eversmann.”

5. Theobaldia (Theobaldia) subochrea, Edw.
Theobaldia annulata var. subochrea, Edwards in Wesenberg-Lund, Danske Vid.
Selsk. Skr., Nat. Math. Afd. (8) vii, p. 198 (1921).

Theobaldia subochrea, Edwards, Ent. Tidsk. p. 50 (1921).
? Culex penetrans, Robineau-Desvoidy, Mem. Soc. Hist. Nat. Paris, iii, p. 407
(1828).

Though structurally identical with T. annulata, this differs so conspicuously and
sharply in coloration that it must be regarded as a distinct species, especially as it
is not confined (as was at first thought) to desert areas, and its coloration therefore
cannot be purely adaptive. The differences from 7. annulata are as follows :—

Mesonotum with the integument lighter, the scales almost uniformly reddish brown.
Abdomen almost uniformly ochreous, the dark brown scales of T. annulata being
replaced by light ochreous brown, and the white ones by almost the same colour.
Whitish lateral patches, however, remain at the base of each segment, and the basal
segmental bands, as well as the median line of the second segment, can be faintly
made out on account of their slightly lighter colour. Leg-markings as in T. annulata,
but somewhat less conspicuous owing to the dark parts being somewhat lighter.
Wing-spots very faint, the scales being less densely aggregated than in T. annulata ;
in the male the spotting of the wing is scarcely perceptible at all.

The larva, so far as I can see, is absolutely identical with that of T. annulata.

Robineau-Desvoidy’s description of C. penetrans applies in many respects to this
species, but he states that the third joint of the middle tarsi of the male is bristly,
which if true, is very remarkable. The type being lost, C. penetrans may be left
as a possible synonym of 7. subochrea.

Distribution —I have examined specimens from the following places :—Meso-
potamia (Basra, Capt. P. J. Barraud; a series including the type male) ; Persia
(Ghilan, 950m., Calhors, J. de Morgan); Palestine (Jerusalem, Dr. Goldberg) ;
Macedonia (Hadji Geul, Capt. J. Waterston) ; Denmark (brackish-water swamp
near Copenhagen, Dr. C. Wesenberg-Lund) ; England (Earl’s Court, London, W. J.
Pendlebury). In Mesopotamia this was the only form found ; in the other countries
T. annulata was found in the same localities.

Subgenus Gulicella, Felt.

6. Theobaldia (Culicella) morsitans (Theobald).

Culex morsitans, Theobald, Mon. Cul. ii, p. 8 (1901).
? Culex flavirostris, Meigen, Syst. Beschr. vi, p. 242 (1830).

Apart from the characters mentioned in the key, this species can generally be
distinguished from T. fumipennis by its slightly smaller size, by the more slender
and rather less hairy male palpi, and by the absence of distinct A-shaped black
marks on the abdominal sternites in both sexes. I at one time considered that the
North American T. dyari (Coq.) might be synonymous, but Dr. H. G. Dyar informs
me that the two are distinct by hypopygial characters.

Meigen’s statements concerning the proboscis and palpi of his C. flavivostris
(male) might possibly be taken as indicating this species, but he says “ Fiisse ganz
braun,” which presumably excludes it. Like the great majority of Meigen’s names,
C. flavirostris is unrecognisable.

The larva varies in colour, but is usually dark brown or blackish.

290 F. W. EDWARDS.

Distribution —Throughout Europe, from Britain and France to Finland (Sam-
matti, Sahlberg; Kuusté, Lundstrém), Petrograd (3 in coll. B. Lichtwardt) and
Macedonia. Not yet known from North Africa or Asia.

7. Theobaldia (Culicella) fumipennis (Steph.) (fig. 5c).

Culex fumipennis, Stephens, Zool. Journ. i, p. 453 (1825).
Culex ficalbii, Noé, Bull. Soc. Ent. Ital. xxxi, p. 238 (1899).
Culicada theobaldi, de Meijere, Tijd. v. Ent. liv, p. 142 (1911).

Distinguished by the characters mentioned in the keys and under T. morsitans.

The larva, apart from the conspicuous structural differences in the siphon, may
generally (perhaps always) be distinguished in life from that of T. morsitans by its
pale yellowish green colour. Both species are mainly bottom feeders and are generally
found in shallow, weedy, stagnant water.

Distribution.—Throughout Europe, except perhaps in the far north. The
material I have examined shows a range from Scotland and France to Sweden and
Macedonia.

Genus Orthopodomyia, Theo.

This genus is well characterised in the larval state by the absence of a pecten
on the siphon and the development, in the fourth-stage larva, of dorsal chitinous
plates on the sixth, seventh and eighth segments of the abdomen. These two
characters together will distinguish the genus from all other mosquitos. In addition
there are some small peculiarities, such as the development of reddish pigment in
the body of the larva, and the very long single lateral hairs on the thorax and
abdomen. The adult characters are not very well marked, the most obvious being :
(1) the presence of only two pro-epimeral bristles ; (2) the small number of bristles
on the pre-alar prominence of the pleurae ; there are five or six in O. pulchripalpis,
but the number is reduced to one or none in the tropical species, while in Culex,
Aédes, Theobaldia and. Taeniorhynchus there are 12 or more; (3) the length of the
first front tarsal joint, which in both sexes is distinctly longer than the remaining
four together, while in most other mosquitos it is only about as long ; (4) the rather
long and stout antennae of the male, all the joints being longer than usual ; (5) the
short fourth joint of the front and middle tarsi of the female, which is much shorter
than the fifth, and like that of the male, scarcely any longer than broad. The very
long fork-cells and the long first hind tarsal joint are also noticeable. The hypopygial
structure is practically the same as in Theobaldia (subgenus Culicella).

1. Orthopodomyia pulchripalpis (Rondani).

Culex pulchripalpis, Rondani, Bull. Soc. Ent. Ital. iv, p. 31 (1872).
Orthopodomyia albionensis, MacGregor, Journ. R.A.M.C. p. 401 (1919);

The beautiful pattern of narrow, pure white lines on the deep black thorax is not
to be seen in any other species known from the Palaearctic region, though confusion
might be possible with some Himalayan species of Aédes, such as A. pseudotaentatus.
The entirely white terminal joint of the male palpi, and the rather long female palpi
(nearly half as long as the proboscis), will distinguish it from all such species, without
reference to the generic characters. The North American O. signtfer (Coq.) is very
closely allied, being structurally identical, but differing in having scattered white
scales on the wings, especially a patch of white scales in the middle of the wing over
the cross-veins. The Oriental species are very differently coloured.

In Ficalbi’s redescription of Rondani’s types he does not definitely describe the
thorax, but refers to the mesonotal scales as “ giallo-ottono?”’ I therefore at first
considered that the British and Italian forms could not be the same. However,

MOSQUITOS OF THE PALAEARCTIC REGION. 291

Prof. M. Bezzi has kindly re-examined for me Rondani’s types in the Florence Museum,
and reports that though the thorax in both specimens is much damaged by the pin,
it is black, ‘‘ even deep black,’’ and there are traces of white lines, and two white
spots in the middle of the scutellum. There can therefore be little or no doubt of
the identity of C. pulchripalpis and O. albionensis.

The larva lives in tree-holes, and shows the modifications commonly found in
such a habitat. It has been described by MacGregor and Lang.

Distribution.—Italy ; England (Epping Forest); France (near Paris, Séguy).
The species may not be so rare as is supposed, since all the specimens so far known
(with the probable exception of Rondani’s types) have been obtained by rearing.

Genus Taeniorhynchus, Arrib.

This genus may be distinguished in the adult from Culex by the absence of
empodia ; from Theobaldia by the absence of spiracular bristles; and from Aédes
by the absence of a definite “tibial scraper’’ (a close-set row of bristles at the tip),
by the non-retractile eighth segment of the female abdomen and the structure of
the male hypopygium. The wing-scales vary greatly in width in the different species,
but are nearly always broader than in Aédes or Culex. Although no more satis-
factory distinctions can be discovered in the adults, the larval siphon is so wonder-
fully modified that on this character the genus is extremely well marked. At the
same time the larval structure of AMfansoniodes is so nearly identical with that of
Taeniorhynchus and Coquillettidia that the three are much best regarded as subgenera
under the main genus Taeniorhynchus, in spite of some rather conspicuous differences
in the adults.

The only known Palaearctic species* belongs to the subgenus Coquillettidia,
to which belong also the North American species and the rather numerous African
and Oriental species placed by Theobald in Chrysoconops. Coquillettidia differs
from the other subgenera in having no post-spiracular bristles, and this affords
a further distinction from Aédes, in which a few of the bristles are apparently
always present.

Taeniorhynchus (Coquillettidia) richiardii (I*ic.).

Culex richiardii, Ficalbi, Bull. Soc. Ent. Ital. xxi, p. 50 (1889), and xxxi, p. 199
(1899).

The rather broad wing-scales and the pale ring in the middle of the first joint

of the hind tarsus will distinguish this species from other mosquitos in the European
fauna.

The bionomics and morphology of the early stages have been described in detail
by Wesenberg-Lund.

Distribution Apparently occurs somewhat locally throughout Europe, except
probably in the far north, and extends into Palestine. Some apparently new records
are: Sweden (Smaland, Stockholm, Boheman); Hungary (Keszthely, Kertész) ;
Austria (Freistadt, Frauenfeld).

Genus Aédes, Mg.

This genus, as a whole, is characterised as follows :—Proboscis of uniform
thickness throughout. Palpi of the female less than one-quarter as long as
the proboscis. Antennae distinctly plumose in the male, with the last two joints
elongate ; with moderately long verticils in the female, all the flagellar joints
being about equal in length. Eyes distinctly separated. A continuous row of
orbital bristles. Pronotal lobes widely separated. Pro-epimeral bristles about

* The South American T. titillans has been recorded from Rumania by Leon, but no doubt
incorrectly.

292 F. W. EDWARDS.

4-6, in a posterior row overlapping the spiracle. Spiracular bristles absent.
Post-spiracular, pre-alar, sternopleural and upper mesepimeral bristles all present
and generally numerous. Postnotum without setae. Eighth segment of female
abdomen retractile, a wide membrane between it and the seventh. Side-pieces
of male hypopygium with a lacuna of chitinisation extending the whole length
of the inner side; claspers articulating in an horizontal plane. Tenth segment
with tergites feebly developed; sternites simple, without teeth or spines. Hind
tibiae with the usual row of fine microscopic hairs just before the tip on the
inner side, and also with a row of 7-10 longer hairs parallel with the first row
and slightly more distally placed. First hind tarsal joint shorter than the
tibia. Pulvilli absent. Front and middle claws of female nearly always
toothed. Cell R, (upper fork-cell) seldom much longer than its stem. Vein A,
(sixth longitudinal) terminating distinctly beyond the level of the base of R, (second
vein). Distinct microtrichia on wing-membrane.

Fig. 5. Structural details of Culicidae: a, tip of hind tibia, seen from the inner side, of Culex

hortensis (scales omitted), x 65; b, the same structure in A édes lepidonotus ; c, the same in Thectaldia

fumipennis ; d, tip of last tarsal joint of Lutzia vorax, showing claws, empodium and pulvilli,

x 340; e, tip of last tarsal joint of Theobaldia annulata, showing claws, small empodium, and

absence of pulvilli, x 340; f, tip of abdomen of dAédes lepidonotus 9, showing slender eighth

segment with narrow sternite, x40; g, the same, Aédes geniculatus 0, showing stouter eighth
segment with broader sternite.

Larva.—Mouth-parts not specially modified for predaceous habits, but the inner
hairs of the mouth-brushes are generally more or less serrate. Antennal tuft generally
at or before the middle. Abdomen without chitinous plates except for the anal
saddle, and sometimes small plates at the bases of the thoracic hairs. Eighth segment
with a lateral comb or patch of scales. Siphon unmodified, short and stout, at most
four times as long as its breadth at the base, provided with a well-developed pecten
and a single pair of ventral hair-tufts, situated about or beyond the middle ; only
very exceptionally with accessory dorsal hairs or hair-tufts.

As pointed out by Dyar (Insecutor Inscitiae, vi, p. 71, 1918) the genus can be
divided into two main groups, on the presence or absence of distinct claspettes in
the male hypopygium (the term claspette in this connection signifying a definite
structure separated from the base of the upper flap of the side-piece, and terminating
in a flattened appendage or a more or less modified bristle). Dyar’s two groups may

MOSQUITOS OF THE PALAEARCTIC REGION. 293

also be defined on the structure of the mesosome of the aedoeagus. In the group
which possesses claspettes the mesosome is a simple tube, rather lightly chitinised
below and at the sides, membranous above. In the other group the mesosome is a
paired structure, the halves of which are more or less crenulate, spiny or brush-like.
Both these distinctions are so well marked that one would expect to find corresponding
distinctions in the body characters or in the larvae, but all efforts in this direction
have so far been fruitless.

The first group includes the subgenera Ochlerotatus and Finlaya, and is spoken of
by Dyar as the New World type—inappropriately, since it includes the majority
of the European mosquitos, and has many representatives in Australia, while Pinlaya
attains its strongest development in the Oriental region. The second or Old World
group (subgenera Ecculex, Aédes and Stegomyia) is almost confined to the tropical
and subtropical regions of the Old World.

The subgenera occurring in the Palaearctic region may be separated (as adults)
by the following keys, which will not necessarily hold good for the species of other
regions.

Male Hypopygium.

1. Claspettes present ; mesosome an unpaired simple structure ae ant ae
Claspettes absent ; mesosome a paired, more complicated structure SD wort
2. Side-piece with more or less distinct apical and basal lobes . . Ochlerotatus.
Side-piece without apical or basal lobes vs as es Finlaya.
3. Clasper deeply divided, placed before tip of side-piece ; without articu-
lated spine... a a8 es - ae: a a Aédes.
Clasper not divided, placed at tip of side-piece, with distinct articu-
lated spine... a = oe of til bys 8 Seed
4. Spine of clasper well before the tip... shi ia or .. Ecculex.
Spine of clasper at the tip (except in A. vittatus) .. one .. Stegomyia.

Other Adult Structures.
1. Proboscis distinctly longer than front femora; last two joints of male

palpi distinctly swollen, hairy, and turned downwards .. ‘i see Ree
Proboscis not longer than front femora; male palpi otherwise ; female
Gere? SHOLt 7s: oP sei es Sa - be 3% si aaa
2. Female cerci short, the eighth sternite large and prominent in repose ;
male palpi somewhat shorter than the proboscis . . ce .. Finlaya.
Female cerci long, the eighth sternite much smaller and not prominent
in repose ae ie He oe ; e ae eae ae
3. Male palpi longer than the proboscis .. se ae Ay Ochlerotatus.
Male palpi not longer than the proboscis is - ae a2 Eecoulex.
A. Palpi alike in the two sexes, very short ; tarsi dark.. ae Ss Aédes.
Male palpi long, slender, the last two joints upturned and nearly bare ;
tarsi with white rings at the bases of the joints .. “F .. Stegomyia.

The adults of these subgenera are treated separately, but it will be more convenient
to tabulate all the known larvae together, as follows :—

Key to the known Larvae of Palaearctic Species of Aédes (sens. lat.).*

1. Antennal tuft represented by a single minute hair; shaft devoid of
small spines ; eighth abdominal segment with a well-marked comb of

teeth set inasingle row... : 2

* This key will not necessarily apply to any but fourth-stage larvae. For the characters
of A. dorsalis and A. intrudens I have relied on American descriptions, and for those of A. alpinus
on Wesenberg-Lund’s description of specimens from Greenland,

10.

F. W. EDWARDS.

Antennal tuft at least two-haired; shaft nearly always spinose ;
eighth abdominal segment with a patch of scales or teeth which ey

tend to form a single row when few in number .. by “f 3 5
. Antennae rather long; dorsal surface of abdomen with numerous
stellate tufts (Finlaya) oe tke aids
Antennae short; dorsal surface of abdomen ‘without conspicuous
stellate tufts: (Ste gomyia) .. 2s “ es ftv HA
. Abdominal tufts composed of long, stort phen x os echinus, Edw.

Abdominal tufts composed of shorter and much more slender bristles
geniculatus, Oliv.

. Comb-teeth simple m3 es .. albopictus, Skuse.
Comb-teeth with lateral ‘basal denticles as .. argenteus, Poiret.
. Shaft of antenna quite bare, tuft 2-3- haired (Seaemina .. vittatus, Bigot.

Shaft of antenna spinose (very slightly so in mariae), tuft generally multiple 6

Fig. 6. Heads of larvae: a, Aédes echinus, Edw.; b, Aédes mariae (Serg.).

. Pecten with detached, simple teeth outwardly ; hair-tuft very small and

well beyond middle of siphon; no long hairs near middle of front

margin of thorax .. 7
Either the pecten has no detached simple teeth outwardly, ot the hair-

tuft is larger and situated practically at the middle of the siphon ;

long hair-tufts present near middle of front margin of thorax

(Ochlerotatus) .. Pale les
The three post-antennal aes een nese in a Siren tie (Aédes)

cinereus, Mg.
Middle post-antennal hairs almost directly in front of the inner (Ecczulex)

vexans, Mg.

. Siphon with three or four pairs of hairs on dorsal (anterior) surface
bio Rossi.
Siphon without hairs on dorsal surface si - 9
. Antennae longer than the head bs AN oP dihentarnsh H. D. & K.
Antennae shorter than the head a os oe : ee
Siphonal index about 1-5 a ae ae sire a, us he SEL

Siphonal index 2 or more ; antennal spinules well-developed oa ht EZ

24.

29.

MOSQUITOS OF THE PALAEARCTIC REGION. 295

. Antennae almost devoid of spinules ; pecten-teeth very long mariae, Serg.
Antennal spinules well-developed ; pecten-teeth shorter .. zammittit, Theo.
. Five or six small hair-tufts in anal brush before the barred area .. ie)
At most three such tufts ; siphonal index 3 or less =f S ae i LF

. Siphonal index deen, over 3; terminal teeth of pecten more or
less detached . oe pe > PIA
Siphonal index at most 3; all pecten- teeth close-set | ee ae jeu. LG

. Comb of eighth segment with 6-9 teeth in one irregular row

semicantans, Mart.
Comb with 23-35 teeth in a triangular patch - of PEA lio)

. Siphonal index about 3-2; anal gills half as long as the Sani lutescens, F.

Siphonal index quite 3-5; anal gills as long as the saddle .. excrucians, Walk.

. Siphonal index 2-3-2-7 58 ab a3 ae ay annulipes, Mg:

Siphonal index 2-6-3... ey .. maculatus, Mg-

. One or two simple detached een! eth Shevewal the siphonal tuft

cataphylla, Dyar.
8

No pecten-teeth beyond the tuft sis bys ‘a 54 is ae On
. Anal gills much shorter than the saddle ¥. cP A. ar cate EO
Anal gills at least as long as the saddle sie Si: . is htm Lae
. Hair-tuft distinctly beyond middle of ae anal gills, one-third to
two-thirds as long as saddle ase ot bis caspius, Pall.
Hair-tuft practically at middle of siphon Se ste am ins ies. 20
. Siphonal index 2; anal gills sakes oe z a i detritus, Hal-
Siphonal index 2- (ee st ts sani veo
. Anal gills globular, not a ater as ine as ene Tie ats dorsalis, Mg:
Anal gills half as long as the saddle .. be = .. Salinellus, Edw.
. Comb with 50 or more scales in the patch... sig eas is 6h gia
Comb with 10-20 pointed scales in the patch hs - os | 24

. Comb-scales sharp-pointed ; anal gills twice as long as ine saddle

pullatus, Coq.
Comb-scales blunt-ended, fringed; anal gills about as long as the saddle
communis, De G.

Last few pecten-teeth detached oF a as Bi ia sat Zo
All pecten-teeth close-set o si 2« [26
Anal segment with a complete cheanione mee duty ales as long as the
gills; frontal hairs single .. : alpinus, L.
Anal segment with saddle only, as Tong as the gills : frontal hairs
branched ; .. tntrudens, Dyar,

. Anal segment with a caiipicee anno ring, abbot as long as the gills

punctor, Kirby.
Anal segment with an incomplete ring, only half as long as the gills
sticticus, Mg.

Subgenus Ochlerotatus, Arrib.

Adults. Proboscis slender, distinctly longer than the front femora. Palpi short
in the female ; longer than the proboscis in the male (very rarely of equal length
or slightly shorter), the last two joints and the tip of the long joint swollen and
hairy, the last two joints turned very slightly downwards, the terminal joint generally
a little more slender than the penultimate. Hair-whorls of male antennae rather
irregular, the majority of the hairs projecting either dorsally or ventrally. Vertex
with narrow scales only, the flat ones being confined to the sides of the head. Lower
mesepimeral hairs usually (not always) present. Male hypopygium with long,

296 F. W. EDWARDS.

simple claspers which are provided with a terminal spine; side-pieces with well-
developed basal lobes and more or less distinct apical lobes ; distinct claspettes
always present, terminating in a more or less flattened appendage ; aedoeagus
simple, the mesosome never divided into two halves. Female cerci always well-
developed and long, the eighth segment small, rarely visible at all externally. All
the claws of the female toothed (normally).

Larva. Antennae with a well marked hair-tuft and minute spicules on the
surface. Frontal hairs generally simple or only slightly branched, the anterior pair
immediately in front of the lower. Abdomen without well-marked stellate tufts ;
the scales of the comb of the eighth segment in a triangular patch.

This subgenus includes the great majority of the Palaearctic and Nearctic species
of Aédes, and attains its maximum development in the north temperate parts of
these regions; it is almost or quite unrepresented in the Ethiopian and Oriental
regions, but appears again with numerous representatives in southern Australia,
and one or two in New Zealand. Many species also occur in the Neotropical
region, and it is possible therefore that Australia was colonised by way of South
America, especially as some of the South American, New Zealand and Australian
species show rather marked affinities. We may perhaps assume from this that the
subgenus is an old-established one.

The Palaearctic species are divisible (as adults) into three groups according to
the markings of the tarsi. The first two of these groups appear to be natural
assemblages of species, but the third, with dark tarsi, is less natural and shows more
structural diversity. These groups may be known as the dorsalis-group, the annulipes-
group and the communis-group ; as mentioned below, A. functor, though placed
on account of coloration in the commumnis-group, shows signs of affinity with the
dorsalis-group, while other members of the communis-group (such as A. communis
itself) are probably more nearly related to the annulipes-group. The structure of
the male hypopygium shows the isolated position of A. rusticus and A. lepidonotus,
but suggests an affinity between A. fullatus and the peculiar A. diantaeus.

Adults.

1. Tarsi with pale rings embracing both ends of the joints, the last hind
tarsal joint entirely pale (dorsalis-group) [compare also Finlaya togot|.. 2

Tarsi with pale rings at the bases of the joints only (annulipes-group)
[compare also subgenera Ecculex and Stegomyia, and Finlaya japonica] 7

Tarsi without pale ue oH) Seek also paeeig Aédes

and Finlaya] . oh : on LZ

2. Abdomen with a pale eden dorsal tees in nddition to aneveee
bands ; sometimes entirely pale .. ae. ek
Abdomen with whitish bands at the bases of the segments only . Bit ae

3. Mesonotal scales generally fawn-coloured, with two narrow shite bands

running the whole length; dark and light wing-scales evenly mixed
caspius, Pall.

Mesonotal scales generally duller brown, with two broad, creamy bands in

front ; dark scales aggregated on certain veins .. aes dorsalis, Mg.
4. Wings and legs densely pause with pale scales ; tarsi brown with creamy
Tings | : ee ba si xs
Wings and legs only very slightly speckled : tarsi black with white TANS), 327,16
5. Mesonotal scales uniformly brown or fawn-coloured. . ais mariae, Serg.
Mesonotum with two longitudinal bands of white scales .. zammuttit, Theo.

6. Mesonotal scales bronzy ochreous, not forming distinct markings
pulchnitarsis, Rond.
Mesonotum with whitish median stripe als .. pulchritarsis, Rond., var.

10.
ie
12.

13.
14.

15.

16.

18.

21.

22.

MOSQUITOS OF THE PALAEARCTIC REGION. 297

_ Outer side of hind femora with numerous scattered dark scales ; white

rings of middle joints of hind tarsi less than half as long as the joints ;
thorax dark with some obscure paler markings .. .. maculatus, Mg.

Outer side of hind femora pale except towards the tip; white rings of
middle joints of hind tarsi at least half as long as the joints ..

. Costa at most with scattered pale scales ; abdomen largely dark . 3)

Costa pale-scaled on anterior edge, at ‘east on basal half ; abdominal
scales mainly or all yellow .. : med |

. Abdomen dark, tergites with well- denned Has al are Ce eee in

the middle. .. semicantans, Martini.
Abdomen with ill-defined yellowish bands and scattered yellowish scales.. 10

Mesonotum with a distinct broad median band of dark scales, sides

yellowish ; pale bands of abdomen mainly basal . .. _annulipes, Mg.
Mesonotum with obscure poche pale bands of abdomen largely at apices

of tergites as as - .. excrucians, Wik,
Costa yellow-scaled aly, on ie asa fed - + Ms .. freyt, Sp. 0.
Costa mainly yellow-scaled in its entire length e id . .lutescens, F.

Pro-epimeral scales all flat and straight, the upper ones black .
rusticus, Rossi.

Pro-epimeral scales all ochreous, the upper ones narrow and curved .. 13
Postnotum with a tuft of scales = de ae .. lepidonotus, Edw.
Postnotum without scales - is 2% ca pee
Wing-membrane whitish ; costal scales all pale .. .. albescens, sp. Nn.
Wing-membrane not whitish ; costal scales mostly or all dark .. sty UD
Front and middle femora conspicuously mottled in front with dark and
light scales : $4 ee 3)
Front and middle femora with only a few pale ‘scales in front ss ie

Head and thorax with very dense bristles ; integument deep black. . alpinus, L.
Head and thorax only moderately bristly ; integument of legs paler .. 17

The dark parts of the abdominal tergites with scattered pale scales ; male
palpi usually all dark : 8 detritus, Hal.

The dark parts of the abdominal tergites without scattered pale scales ;
male palpi with a whitish ring or numerous pale scales on the long basal

FOULS 206 Ps de .s - a s% 18
Proboscis all ane : Me is calapnyjilas pyar
Proboscis with numerous pale scales in the middle .. .. salinellus, Edw.

Hind femora pale, with a sharply defined black-scaled area at the tip
diantaeus, H. D. & K.
Dark area at tip of hind femora much less sharply defined .. : 4.
Hind tibia (at least in the female) with a distinct whitish ees on the
outer side running nearly the whole length ; mesonotum with white
scales at the sides, blackish in the middle .. hs sticticus, Mg.
Hind tibia without whitish stripe on outer side in either sex... poemay Al

Scales of head and mesonotum ochreous brown, the mesonotum usually
with one broad dark brown median stripe; sometimes with two
ill-defined stripes, or none... .. punctor, Kirby.

Scales of head and mesonotum darker “brown, often mixed with white at
the sides; the mesonotum ear without distinct stripes, or with

two ill- defined ones .. a3 te he hee
Flat scales of the head soaronchans the qaidate ae ais .. parvulus, Edw:
Flat scales confined to the sides of the head . ; communis, De G. >;

" intrudens, Dyar ; pullatus, Coq:

298

F. W. EDWARDS.

Male Hypopygia.

_ Basal lobe of side-piece bearing a row of flattened bristles or scales ;

terminal spine of clasper wavy es afi BS .. rusticus (Rossi).
Basal lobe of side-piece quite otherwise ; terminal spine of clasper straight 2

. Side-piece with three spines or differentiated bristles ; two on the basal

lobe, backwardly directed, one more distally placed and inwardly directed 3
Side-piece with at most two differentiated bristles on the basal lobe 2)..%,49

. Side-piece swollen in the middle, where it has a large inwardly-directed

hair-tuft - - sis si * ,. diantaeus, H. D. & K.
Side-piece more regularly shaped, without hair-tuft in the middle se, ee

_ A distinct hair-tuft adjoining the apical lobe; stem of claspette with a

distinct projection near the middle. . a nf intrudens, Dyar.
No definite hair-tuft adjoining the apical lobe ; stem of claspette somewhat

angulated but without definite projection. . <0 .. pullatus (Coq.).

Fig. 7.

Claspettes of Palaearctic species of Aédes, dissected from hypopygia and drawn

in side view, all x 75: a, A. maculatus (Mg.); b, A. semicantans, Mart.; c, A. lutescens (F.) ;
d, A. freyi, sp.n.; e, A. annulipes (Mg.); f, A. excrucians (Walk.); g, A. punctor (Ixirby) ;
h, 4. detritus (Hal.); i, A. pullatus (Coq.); j, A. communis (DeG.); k, A. salinellus, Edw. ;

10.

1, A. cataphylla, Dyar.

. Basal and apical lobes of side-piece both very small, the latter hardly

distinguishable ns ie ate we _ fd
Either basal or apical lobes, or both, quite large and conspicuous 5-3

. Appendage of claspette short and twisted, rounded at tip lepidonotus, Edw.

Appendage of claspette long, curved and pointed .. Ay

. Basal lobe without a spine re Be mariae (Serg.) ; zammittii (Theo.).
Basal lobe with a distinct spine zl HH. pulchritarsis (Rond.).
. Basal lobe with two spines ; apical lobe ill-defined .. ae ae ee
Basal lobe with at most one spine ; apical lobe nearly always well-marked.. 10
. Basal lobe very prominent (fig. 8b) .. Pe 7 es dorsalis (Mg.).
Basal lobe much less prominent (fig. 8 a) bk a .. casptus (Pall.).

Apical lobe reaching back to near middle of side-piece, and clothed with
very short, curved bristles; stem of claspette short and straight .. 11
Apical lobe not reaching so far back, and bearing longer bristles sigh

MOSQUITOS OF THE PALAEARCTIC REGION. 299

11. Appendage of claspette three times as long as broad; basal lobe not

separated from the side-piece .. punctor (Kirby).
Appendage of claspette not much longer than broad ; basal lobe separated
from the side-piece in its apical portion 26 uy sticticus (Mg.).
12. Basal lobe with a strong black spine and short beste as lutescens (F.).
Spine of basal lobe when present weaker and ae and eran ia accom-
panied by long bristles ba, : 5 AS
13. Basal lobe small, without spine, but a rugose area ees more Wee hall
the length of the side-piece .. . 14
Basal lobe larger, spine generally well- marked, rugose area not near ly
reaching middle of side-piece (except in frey?) .. ee 15
14. Stem of claspette uniformly stout de ms ays a Pie (Mg,).
Stem of claspette slender apically .. es a excrucians (Walk.).
15. Stem of claspette shorter and nearly straight ae - ss 1480
Stem of claspette longer and strongly curved : fs asa q
16. Appendage of claspette winged in the middle Sale. are on the
basal third; basal lobe rather small, not pointed .. detritus (Hal.).
Appendage of claspette winged for its whole length or nearly ; basal lobe
produced and rather sharply pointed ois 17
17. Basal lobe much longer than broad ; aE a Pesce nee
broad .. .. maculatus (Mg.).

Basal lobe scarcely longer than broad ; appendage of claspette less broad
semicantans, Mart.

18. Side-piece with short hair only Bs i jay OS
Side-piece with long hair arching over ‘the upper surface .. oP “a 2
19. Basal lobe large, spine present though weak i ee freyt, sp. n.
Basal lobe smaller, spine absent ay .. parvulus, sp. n.

20. Appendage of claspette with two slight Hdeesn near ihe base communis (De G.).
Appendage of claspette with one ridge or wing, which is broadest in the

middle .. be s3 dis a6 Si ak
21. Aedoeagus and anal seenieat fa givemnised ‘ eee (Lo).
Aedoeagus and anal segment not unusually strongly chitinised #s 3 2,

22. Lobes of ninth tergite with about six short straight bristles .. cataphylla, ea:
Lobes of ninth tergite with about twelve longer bristles which curve
outwards Zt i #4 és $a 42 .. Ssalinellus, Edw.

1. Aédes (Ochlerotatus) caspius (Pallas) (= dorsalis, Theo.) (fig. 8 a).

Culex caspius, Pallas, Reise versch. Prov. Russ. Reichs. i, p. 475 (1771).

Culex punctatus, Meigen, Klass, i, p. 6 (1804). .

? Culex siculus, Robineau-Desvoidy, Mém. Soc. d’Hist. Nat. ii, p. 406 (1827).

Culex penicillaris, Rondani, Bull. Soc. Ent. Ital. iv, p. 31 (1872).

? Culex leucogrammus, Loew, Zeitschr. Ges. Naturw. xliii, p. 413 (1874).

Grabhamia subtilis, Ed. & Et. Sergent, Bull. Mus. Paris, xi, p. 240 (1905).

Grabhamia willcocksi, Theobald, Mon. Cul. iv, p. 294 (1907).

Grabhamia longisquamosa, Theobald, Ann. Mus. Nat. Hung. ui, p. 102 (1905).

Mansonia arabica, Giles, J. Trop. Med. p. 130 (1906).

Culex arabicus, Becker, Denkschr. k. Ak. Wiss. Wein. Ixxi, p. 140 (1910).

As has often been pointed out, this is a very variable species, particularly as
regards the abdominal markings. The palest specimens have the abdominal scales
uniformly ochreous, while in the darkest the median pale stripe tends to be reduced,
one Hungarian specimen which I have examined showing it on the second segment
only. A. caspius may generally be known by the bright fawn-coloured scales of the
mesonotum, with two narrow longitudinal stripes of white running the whole length,
and by the even scattering of the dark scales over the wing-veins, as well as by the

300 F. W. EDWARDS.

shape of the basal lobes of the side-pieces of the male hypopygium. The variation
in the thorax is in the direction of the reduction of the white scales, which are
occasionally absent, especially in the male sex, where the reduction of the white lines
is often accompanied by a more or less pronounced bleaching of all the mesonotal
scales. There seems to be a slight difference between the north and south European
forms in coloration, the Mediterranean type being lighter than the north European,
and also somewhat smaller. The difference, however, is quite indefinite, and I see
no necessity for the maintenance of a varietal name. Theobald’s G. longisquamosa
was evidently described from an extremely pale specimen. The variety hargreavesi,
Edwards (Bull. Ent. Res. x, p. 130, 1920), seems to be well distinguished by having
all the scales of the female mesonotum white; it is, however, known from only a
very few rather imperfect specimens.

There are two main generations in the year (in June and September), and the
species is often abundant enough to cause serious annoyance. It will often migrate
some miles from its breeding-places (a habit common to most salt-marsh species),
and is the only Aédes in the European fauna which commonly enters houses. The
adults are not known to hibernate, but Dr. Langeron captured a female at Gabes in
January 1919.

The larvae show the reduction in the size of the anal gills which is commonly
seen in salt and brackish-water species, but they are by no means confined to saline
waters, the species often spreading far up the courses of the larger rivers, breeding
in open meadows. It may be that specimens from fresher water have longer gills,
as these organs certainly vary in length in this species. English specimens which
I have examined agree with Martini’s description in having the gills about two-thirds
as long as the saddle, while Capt. Barraud’s Mesopotamian specimens agree with
Wesenberg-Lund’s description in having gills only about one-third as long as the
saddle.

Distribution.—European and Mediterranean coasts ; Central European plain, and
eastwards to the Gobi desert ; desert regions of North Africa; Palestine; Persian
Gulf; Punjab, as far inland as Rawalpindi. Apparently does not extend into
North America. Since the species has been so much confused with A. dorsalis, it
may be worth while to record some of the specimens I have examined from different
museums: Denmark (near Copenhagen, Wesenberg-Lund ; Finland (Kuusté,
Lundstrém) ; France (Bourg la Reine, Langeron); Germany (Usedom, Sulldorf,
Litchwardt coll.) ; Austria (Vienna, Handlirsch; Carniola, Lowtsch) ; Hungary
(Neusiedler See, Handlirsch ; Hortobagy, Kertész; Csepel, Bartko; Fehertelep,
Ujhelyi ; Iszak, Uhl; Torda, Bird); Italy (Mehadia, Livorno, Spalato, Mann) ;
Roumania (Tultscha, Mann); Constantinople (Paris Mus.); Asia Minor
(Fregli, Tskehir, Kara, Lendl); Palestine (Jerusalem, Goldberg); West Caspian
(‘“Lenkoran bis Elizabetpol, 1-2 Aug. Nachts. Schrecktl. Miicken-plage,” Berlin
Mus.) ; Transcaspia (Aschabad, Firudza, Tedjin and Kopet Dagh, C. Anger) ; Gobi
Desert (Cha Tcheou, Marais de Pa-hou-lian, Dr. L. Vaillant).

2. Aédes (Ochlerotatus) dorsalis (Meigen) (ec Theobald et al.) (fig. 8 b).

Culex dorsalis, Meigen, Syst. Beschr. vi, p. 242 (1830).

Culex maculiventris, Macquart, Dipt. Exot. Supp. i, p. 7 (1846).
Culex curriei, Coquillett, Can. Ent. xxxili, p. 259 (1901).

Culex onondagensis, Felt, N.Y. State Mus. Bull. 79, p. 278 (1904).
Grabhamia broquettii, Theobald, Entomologist, xlvi, p. 179 (1913).
Aédes grahami, Ludlow, Insecutor Inscitiae, vii, p. 154 (1920).

Doubt has been expressed by Wesenberg-Lund whether this is really distinct
from A. caspius, but I still believe that the characters I have adduced are sufficient
for the separation of the two, though it must be admitted that they are both variable
species with entirely similar habits, and that they are frequently found together.

MOSQUITOS OF THE PALAEARCTIC REGION. 301

The thoracic scaling (brown to dark brown central stripe on front part of mesonotum
only, with a broad creamy stripe on each side of it, dark brown again on the shoulders),
wing scaling (preponderance of dark scales on the first, third and fifth veins, and of
pale scales elsewhere), and the prominent basal lobes of the side-pieces of the male
hypopygium will render the identification of nearly every good specimen certain.

I have decided to adopt the name dorsalis for this species for two reasons. First,
I am indebted to M. Séguy for sending me a female from Meigen’s collection labelled
“Culex dorsalis. Berlin,’’ probably in Meigen’s own handwriting. As C. dorsalis
was originally described from Berlin, it seems reasonable to accept this specimen as
the actual type of the species. It is in good condition, and obviously A. curriet,
not A. caspius. Secondly, from the descriptions of Staeger and Zetterstedt, as
well as from some examples named by Staeger which were sent me by Dr. Wesenberg-
Lund, it would seem that these writers based their conception of C. dorsalis mainly
on A. curries.

Fig. 8. Hypopygia of Palaearctic species of Aédes: a, outline of side-piece, to show structure

of basal lobe, of A. caspius (Pall.) ; b, the same, A. dorsalis (Mg.) ; c, the same, A. mariae (Serg.) ;

d, hypopygium of 4. freyi, sp. n.; e, hypopygium of A. lepidonotus, Edw., ventral view, claspette
shown also in side view; f, the same, 4. parvulus, sp.n.; g, A. intrudens, Dyar.

Rather contrary to expectation, Macquart’s C. maculiventris proves to be this
species and not A. caspius, according to the type female, which was sent me by
M. Séguy.

The larva is at present only known from the description of Howard, Dyar and
Knab ; it has not yet been isolated in European collections. Specimens sent me by
Dr. Dyar agree with the description in the monograph, and differ from A. caspius
in at least three points: the siphon is distinctly more slender (index about 2°75
instead of about 2-3); the hair-tuft is scarcely beyond the middle of the siphon ;

(4183) y

302 F. W. EDWARDS.

and the anal gills are very small and globular, not a quarter as long as the saddle.
Whether these characters distinguish A. dorsalis as a species or merely the American
race remains to be proved.

Distribution.—Coasts of northern Europe from France and southern England
to Norway and the Baltic ; Central European plain, and thence westward across
Central Asia, China and North America as far as the Atlantic coast ; North Africa
(according to Macquart).

Some localities from which |] have examined specimens are :—Denmark (Wesen-
berg-Lund) ; Sweden (Ostergétland, Haglund); Finland (Kexholm, Eriksberg,
Uskela, E. J. Bonsd.; Kuusté, Lundstrom; Soroka, J. Sahlberg); Germany
(Berlin, Meigen, Enderlein ; Usedom, Siilldorf, Lichtwardt; Halle, Loew); Austria
(Hainfeld, Mzk); Hungary (Budapest, Kertész ; Budafok, Bartko ; Fehertelep,
Ujhelyi ; Yorda, Bird; Keczel, Hild, Thalhammer ; Neusiedler-See, Handlirsch) 3
North Russia (Waloniki, Velitchkovsky) ; Mongolia (“ Vallée prés de la Kouré de
Bandie, 1500 m.’’ and ‘‘ Bords du Tarim,’ Mission de Lacoste, Dr. du Chazaud,
1909) ; Siberia (Irkutsk, Schulz); North China (Tinghai, C. Ford).

Fig 9. Aédes mariae (Serg.), end of abdomen of larva; comb and pecten teeth and mentum
more highly magnified.

MOSQUITOS OF THE PALAEARCTIC REGION. 303

3. Aédes (Ochlerotatus) mariae (Sergent) (figs. 6 b, 8c, 9).

Culex mariae, Ed. & Et. Sergent, Ann. Inst. Pasteur, xvii, p. 62 (early in 1903).

Before having examined specimens of the true A. pulchritarsis, | took A. mariae
to be synonymous with Rondani’s species, but the two are in fact distinct. 0.
mariae has the wings, femora and tibiae, and even the tarsi in part, densely speckled
with pale scales ; the tarsal rings are creamy white and not very sharply marked ;
the mesonotal scales are almost “uniformly ochreous without any bronzy tint. The
male hypopygium resembles that of O. caspius, but the basal lobes of the side-pieces
are smaller and carry no spines.

The Sergents’ description of the larva pene somewhat inadequate, fresh figures *
are given here, prepared from specimens collected by Capt. Barraud. These figur es
will explain themselves, but it may be specially noted that the antennae are almost
devoid of small spicules, a very unusual character for this genus ; the siphon is also
remarkable in being almost as broad at the tip as at the base, the base being feebly
or not at all chitinised (even in full-grown larvae) ; siphonal index 1-4-1-6. (It
may be remarked here that Wesenberg-Lund’s figure of the siphon of A. caspius
is inaccurate ; the tip in that species is really much narrower than the base.) The
anal gills in A. martae are minute and globular, as usual in salt-water breeders ; the
saddle is extremely small ; formula of anal brush 10 ++ 4.

The species seems to be exclusively a salt-water breeder.

Distribution.—Algerian coast (Sergent); Southern France (Séguy); Palestine
coast (Athlit, Barraud) ; Syria (Beirut, Barraud). Probably occurs also along the
intervening Mediterranean coasts.

4. Aédes (Ochlerotatus) zammittii (Theobald).
Acartomyia zammuitiii, Theobald, Mon. Cul. iii, p. 252 (25th July 1903).

This rather closely resembles A. mariae, the only difference I can detect in the
adult being the presence of two more or less definite longitudinal stripes of white
scales on the mesonotum, resembling those seen in O. casprus. The male hypopygia
appear to be identical. |The main reason for keeping the two distinct is the occurrence
of certain larval differences. The two larvae of A. zammuttit in the British Museum
are both very much damaged, and one is immature, but the antenna has distinct
spicules, and the pecten teeth are shorter, rather more numerous, closer together,
and with more serrations than in A. mariae ; it is possible they may have been wr ongly
associated with the adults, and in any case it is very desirable that more and better
material should be obtained. If A. zammittit should prove identical with A. mariae,
the latter name will stand for the species ; it must have been published earlier, since
Theobald quotes it (Mon. Cul. ui, p. 354).

Distribution.—Originally described by Theobald from Malta. I have also seen
specimens with a more or less distinctly white-striped thorax from Palma, Majorca
(Grinberg), and Southern France (Dollfus, per Dr. Langeron).

5. Aédes (Ochierotatus) pulchritarsis (Kondani).

Culex pulchritarsis, Rondani, Bull. Soc. Ent. Ital. iv, p. 31 (1872),
Culex leucacanthus, Loew, Beschr. Eur. Dipt. in, p. 1 (1873).

In the specimens which I have examined there are no pale scales on the wings, the
femora and tibiae are moderately speckled ; the tarsi are black, with sharply marked
white rings; the mesonotal scales rather variable, usually dull bronzy-ochreous
with some patches of darker ones, sometimes, as in the specimens from Paris and
Chitral, a more or less distinct central white stripe. It is just possible that the species
may not be correctly identified, since Ficalbi in his detailed description says of the

(4183) ¥2

304 F. W. EDWARDS.

wings, “con qualche brizzolatura chiaro.” Ficalbi’s description, however, fits this
species much better than it does O. mariae. The two Italian specimens were both
determined by Prof. M. Bezzi, who has seen Rondani’s collection. Judging from
the description, it seems highly probable that Loew’s C. leucacanthus is the same
species.

The larva is unknown. The male hypopygium differs from that of A. mariae
and A. zammittii in having a single distinct spine on the basal lobe of the side-piece, .
the structure being otherwise similar.

Distribution.—Italy (Toscana, near sea, Ficalbi; Faenza, E. Hargreaves ;
Macerata, Bezzi); Macedonia (Bajirli, near Snevce, Waterston) ; Persia (Kasan,
Loew); Paris (Berland, Ségwy) ; Croatia (Novi, Horvath) ; Chitral (Drosh, Walker,
1915; per Capi. BP. J. Barraud):

6. Aédes (Ochlerotatus) maculatus (Meigen) (fig. 7 a).

Culex maculatus, Meigen, Klass. und Beschr. i, p. 4 (1804).
Culex cantans, Meigen, Syst. Beschr. i, p. 6 (1818).
Culicada waterhouset, Theobald, Ann. Mag. Nat. Hist. (7), xvi, p. 674 (1905).

This species may generally be distinguished from others of the group by its
darker colour ; the femora (especially the hind pair) have more numerous dark scales,
the white rings on the tarsi are narrower, and the mesonotum is much darker, the
pale scales occurring mainly on the posterior third and often forming a pair of spots
just behind the middle and indistinct pale lines in front of the scutellum. There
appear to be two more or less definite forms: a smaller, with entirely dark-scaled
wings and distinct pale bands at the bases of the abdominal tergites, and a larger,
with scattered pale scales on the wings and the abdominal bands more or less obsolete,
the pale scales which are present occurring as much towards the apices as the bases
of the segments. The peculiar male hypopygium, with its divided basal lobes and
very broad appendage to the claspettes, is the same in both forms.

I am still unable to distinguish satisfactorily the larvae of A. maculatus and
A. annulipes,* but the two species are certainly quite distinct. A. maculatus is
essentially a woodland species, while A. annulipes (and most others of the group)
prefer more open breeding-places.

This species has generally been known as A. cantans, though Meigen’s description
will apply as well or better to one of the other species of the group. ‘or this reason
Lang adopted the later name waterhousei. M. Séguy, however, informs me that males
of this species are labelled C. cantans in Meigen’s collection in Paris, and the identi-
fication of Theobald, de Meijere and others must therefore be regarded as correct.
Meigen himself states that his C. maculatus was the male of C. cantans, and this state-
ment must be accepted, since the type of C. maculatus no longer exists to prove or

disprove it. Many of the earlier records of C. cantans apply no doubt to other species
of the group.

Distribution.—Apparently confined to Europe, occurring chiefly north of the
Alps. I have seen males from Britain; Frarice (Ségwy, Langeron); Belgium
(Goetghebuer) ; Denmark (Wesenberg-Lund); Sweden (Stockholm, Boheman ;
Smaland, . Haglund); Finland (Kymmene, Sallm.); Germany (Urdingen and
Frankfort-an-der-Oder, Riedel ; Berlin, Oldenberg, Lichtwardt) ; Austria (Polzleinsdorf,
Schiner ; Moritzburg, Kuntze); Hungary (Munkacs, Uyhelyi); also females,
probably of this species, from Lappland (Qvikkjokk), and Italy (Turin, Sangone,
coll. Bezzt).

* Séguy’s figure of ‘“‘ A. cantans”’ in Bull. Mus. Paris, 1920, p. 327, really represents A.
communis ; his later figure of ‘‘ O. cantans’”’ in Bull. Soc. Ent. France, 1920, p. 310, probably
represents A. lutescens (see Séguy, Bull. Soc. Ent. France, 1921, p. 165).

MOSQUITOS OF THE PALAEARCTIC REGION. 305

7. Aédes (Ochlerotatus) semicantans, Martini (fig. 7 b).

Aédes semicantans, Martini, Arch. f. Schiffs- und Tropenhyg. xxiv, Beiheft 1,
p. 247 (1920) (diagnosis only), and Sitzb. u. Abh. natf. Ges. Rostock, vii,
p. 205 (1920).

Resembles A. maculatus in coloration, being darker than the other four species
of the group; the mesonotum has a more definite dark median stripe than in
A. maculatus, and the abdominal segments have distinct basal white bands, which are
narrowed in the middle, but the two are not easily separated, except by characters
of the male hypopygium and larva, which are perfectly distinct. The hind femora
are largely pale on the outer side, and the tarsal rings are broader than in A. maculatus.
One or two North American species resemble this rather closely, but I have not at
present been able to identify A. semicantans definitely with any North American
form. The larva is very distinct on account of the small number of scales in the
comb of the eighth segment.

Distribution —Northern Europe; apparently widely distributed, but local.
I have seen males from Sweden (Smaland, Boheman) ; Germany (Hamburg, Martin ;
Berlin, Stobbe ; Posen?, Loew); Austria? (coll. Winthem).

8. Aédes (Ochlerotatus) annulipes (Meigen) (fig. 7 e).

Culex annulipes, Meigen, Syst. Beschr. vi,.p. 241 (1830).

Aédes quartus, Martini, Uber Stechmiicken, p. 128 (1920).

Meigen mentions the dark stripe in the: middle of the mesonotum, the banded
abdomen and pale femora, and I therefore have no doubt that the species is correctly
identified. Zetterstedt, Ficalbi and others who have described the entirely yellow
abdomen of the female probably had A. lutescens before them. In this species the
abdomen almost always has distinct yellowish-white bands, which are situated
mainly or entirely at the bases of the segments.

The larva is of the short-siphoned type, apparently identical with that of
A. maculatus (except that the siphon is slightly shorter on the average), but very
distinct from that of A. excrucians. It occurs typically in open swamps.

Distribution.—I have examined male specimens from England (various localities) ;
Holland (Ghent, Theobald) ; Belgium (Wesembeek, Tonnoir) ; Germany (Urdingen,
Riedel; Berlin, Oldenberg) ; Lower Austria (Speising, Mik) ; Hungary (Pressburg,
Mik; Tultscha, Mann). A female from Sweden (Ostergédtland, Haglund) may be
this species, but is perhaps more probably A. excrucians. Martini records it from
the neighbourhood of Hamburg and Dantzig. It may be regarded as the western
European representative of A. excrucians, though the ranges of the two species
must overlap to some extent.

9. Aédes (Ochlerotatus) excrucians (Walker) (fig. 7 f).

Culex excrucians, Walker, Ins. Saund. Dipt. p. 429 (1856).

Culex abfitchii, Felt, Bull. N.Y. State Mus. 79, p. 381 (1904) ; et auct.

(2?) Culicada surcoufi, Theobald, Bull. Mus. Paris, xviii, p. 59 (1912).

Aédes excrucians, Dyar, Insecutor Inscitiae, vii, p. 25 (1919), and viii, p. 109

(1920).

This species is most nearly allied to A. annulipes, but the abdomen is less distinctly
banded, the dark bands being reduced to a more or less diamond-shaped spot on each
segment ; the dark colour is often more extensive in the female. The mesonotum
usually has some obscure markings; rarely a distinct, broad, dark median stripe
as in A. annulipes. The difference of structure in the male claspettes is slight but
well defined. I can see no difference between American and European specimens,
except that in the former the pale markings are cream-coloured rather than yellowish.

A. excrucians is evidently one of the commonest species of the group on the
European continent (it has been described by Martini as A. abfitchit), and it is therefore

306 F. W. EDWARDS.

not surprising that it has frequently been determined either as C. cantans or as
C. annulipes. 1 consider, however, that the evidence of the original descriptions
and of the types in Meigen’s collection in Paris is sufficient to allot these names to
other species.

I learn from M. Séguy that the type male of Culicada surcoufi has been lost,
so that there is now very little prospect of determining Theobald’s species with
certainty. A female which M. Séguy sent me for examination might have been either
this species or A. annulipes. A female in Meigen’s series of C. cantans in the Paris
Museum is almost certainly A. excrucians.

Distribution.—I have examined male specimens from the following countries :—
Finland (Hattula, L.v. Essen; Tvarminno, Messuby and Walkj, R. Frey ; Eriksberg,
E. J. Bonsd.; Karislojo, J. Sahlberg); Sweden (Sédermanland, Awnwillius ;
Ostergdtland, P. Wahlberg); Denmark (Wesenberg- Lund); Germany (Berlin,
Lichtwardt, Oldenberg, Stobbe, Enderlein) ; Austria (many in Winthem’s old collec-
tion without definite data; one male labelled “ rufibarbis, Gtl.”) ; Hungary (Buda,
Bird ; Munkacs, Uyjhelyt; Berecsasz, Kertész) ; Saghalin Island (Paul Labbe,
1902). In addition I have seen many females which are probably this species, but
cannot be determined with absolute certainty, including some from Siberia
(Antsiferovo, 59° 10’, and Turuchansk, 65° 55’, Tryvbom), The species may therefore
be assumed to have a continuous distribution over North Europe and North Asia ;
it is also known to be widely spread in North America. It appears to be absent
from North-western Europe.

10. Aédes (Ochlerotatus) freyi, sp. n. (figs. 7d, 8d).

Scales of head and mesonotum all small and yellow. Abdomen yellow, the
tergites with black apical lateral patches, no median dark line. Male palpi longer
than the proboscis by nearly the length of the last joint. Long joint with a yellow
ring at the base and another beyond the middle ; last two joints pale-scaled at the
base ; hairs long, brown. Proboscis all black. Tarsi with white rings at the bases
of the joints; those on the middle joints of the hind tarsi slightly more than half
of the length of the joints. Front and middle femora speckled in front, yellow
behind ; hind femora mostly yellow, with some black scales towards the tip. Wing-
scales mostly dark, the costa yellow on about the basal third.

Hypopygium: side-pieces rather slender, fully three times as long as their greatest
breadth. Basal lobe prominent, rounded apically, with a small patch of long hair,
and a single long, weak, pale spine. Apical lobes rather large, somewhat pointed,
with a few longish hairs. No arching-hairs on sternal side of side-piece. Claspette
with long curved stem and broadly flattened appendage, the expansion commencing
at the base. Lobes of ninth tergite each with about six short hairs.

This species, though resembling A. /utescens in coloration and A. semicantans
in the structure of its hypopygium, is certainly distinct from both. The few examples
I have seen are all more or less damaged and do not allow a very precise definition
of the species on colour characters. Possibly the costa being yellow towards the
base only instead of for the greater part of its length might distinguish the female
of A. freyi from that of A. lutescens. The only undoubted female of A. freyt
I have seen was considerably denuded ; it had scattered light scales on the wings,
and the proboscis largely pale except towards the base and tip.

Distribution.—Finland (Eriksberg, E. J. Bonsd., type male in Helsingfors
Museum) ; Germany (Berlin-Finkenkrug, 27.v.00, L. Oldenberg, 33, 1g):

11. Aédes (Ochlerotatus) lutescens (Fabricius) (fig. 7c).

(?) Culex flavescens, Miller, Fauna Insectorum Friedrichsdalina, p. 87 (1764).
Culex lutescens, Fabricius, Syst. Ent. p. 800 (1775).
(?) Culex variegatus, Schrank, Enum. Ins. Austr. p. 482 (1781).

MOSQUITOS OF THE PALAEARCTIC REGION. 307

Culex flavescens, Fabricius, Syst. Antl. p. 35 (1805).

(?) Culex bipunctatus, Robineau-Desvoidy, Mém.. Socs d’Hist. Nat. Paris, i
p. 405 (1827).

Culex flavus, Motchulsky, Bull. Soc. Imp. Nat. Moscow, xxxii, pt. 2, p. 503
(1859).

Culex flavescens, Theobald, Mon. Cul. i, p. 410 (1901).

Culex arcanus, Blanchard, Les Moustiques, p. 303 (1904).

Culex fletchert, Coquillett, U.S. Bur. Ent. Tech. Ser. 11, p. 20 (19086).

Aédes cyprius, Ludlow, Insecutor Inscitiae, vii, p. 158 (1920).

Although this species seems to be subject to a good deal of variation, it may
probably always be recognised by the predominantly yellow-scaled costa and the
mainly or entirely yellow-scaled abdomen. In the lightest specimens the proboscis
is yellow-scaled except at the tip, where the scales are black ; the palpi and wings
are almost entirely yellow-scaled; the mesonotal scales are rather light yellow ;
the thoracic integument is reddish, with a black patch in front from which three
black lines extend backwards; the abdominal scales are all yellow in the female,
usually with a median line of dark ones in the male ; the tarsi, though with the usual
white rings at the bases of the joints, have the dark portions largely replaced by yellow,
except at the tips of the joints. In the darkest specimens the proboscis and palpi
are almost entirely black-scaled, the wings are much darker, even the costa having a
considerable sprinkling of dark scales ; the mesonotal scales are rather deep brown,
contrasting strongly with the white scales of the pleurae ; the thoracic integument
is nearly all black ; the female abdomen has rather numerous dark scales at the sides ;
and the tarsi are darker. In some males the thoracic scales are almost white, and
there are traces of a darker median band.

It is possible that the two forms above described may represent distinct varieties,
or even species, but they appear to intergrade, and males associated with both dark
and light females have practically identical hypopygia, the only difference observed
being that some (though not all) of the light specimens had a more or less definite
hook to the membrane of the claspette appendage (see fig. 7c). Apart from this
the hypopygium is well distinguished by the shape of the side-pieces, the very large
basal lobes bearing a strong black spine, and the appendages of the claspettes, which
have no membranous expansion on the basal third.

I at first considered that A. lutescens was identical with the American A. fletchert,
but there are some slight differences: in A. fletchert the apical lobe of the side-
piece of the male hypopygium is smaller, and the mesonotum has a distinct broad
dark central stripe.

This, being a common European species, is much more likely to be Fabricius’
C. lutescens than the rare one which Theobald has redescribed as such. Fabricius
emphasises the yellow costa and the yellow proboscis with a black tip ; he does not
mention the tarsi, the statement that they are dark being due to Meigen, who may
have had another species before him. Martini was of the same opinion regarding
Fabricius’ species, but did not adopt the name ; I consider the identification sufficiently
probable for the name to be used. Zetterstedt, Ficalbi and Martini use the name
annulipes, Mg., for it, but this is certainly wrong, since the abdomen is never banded,
as Meigen states is the case in annulipes. Although Motchulsky’s diagnosis is very
brief, I consider it highly probable that his C. flavus is our A. lutescens.

Distribution Europe, except the west; Siberia; extending in a slightly
modified form into North America. I have seen specimens from Denmark (Wesenberg-
Lund) ; Sweden (Ostergétland, Scania, Boheman, light females) ; Finland (Kuusté,
Lundstrém ; Eriksberg, E. J. Bonsd.); Germany (Berlin, Wanen, Lichtwardt,
Oldenberg, Tetens, females of light and dark forms; Radoj, Loew, dark female) ;
Austria (Médling, Pokorny; Michelstettin, Bischoff; Sterize, Styria, Mann ;
Weyshi, Steiermark, Mann ; males, and females of dark form) ; Hungary (Pressburg,

308 F. W. EDWARDS.

Mik; Jaszenova, Ujhelyi ; Kovakspatak, Kertész ; Budapest, Cstki, males ;
Neusiedler See, Mik, and Munkacs, Uyhelyi, dark females) ; Ural (Winthem, light
female) ; Siberia (Nasimovo, 59° 35’, males and dark females; Nikulina, 60° 25’, lighter
male, Tvybom ; Omsk, Grand, light females) ; Kamtchatka (Bolsherjetsk, 20.vii.17,
Y. Wuorentaus, light female) ; Asia Minor (Seraj-Koj, Naday, dark females).

12. Aédes (Ochlerotatus) rusticus (Rossi).

Culex rusticus, Rossi, Fauna Etrusca, ii, p. 333 (1790).

? Culex musicus, Leach, Zool. Journ. ii, p. 293 (1825).

Culex pungens, Robineau-Desvoidy, Mem. Soc. Hist. Nat. Paris, ili, p. 407 (1827).

Culex quadratimaculatus, Macquart, Suites a Buffon, i, p. 34 (1834).

Culex diversus, Theobald, Mon. Cul. ii, p. 73 (1901).

Culex nemorosus var. luteovittatus, Theobald, Mon. Cul. ii, p. 85 (1901).

This is the largest species of the dark-footed group of the subgenus, and is very
distinct from all the rest in the structure of the hypopygium and larva. The male
has the palpi stouter than in most species; it can generally be distinguished even
by the naked eye on account of the dense golden-yellow hairs which arch over the
hypopygium, hiding the claspettes. The most strongly-marked form of the female,
on which I have presumed Rossi’s and Robineau-Desvoidy’s original descriptions
were based, shows a yellowish longitudinal stripe running the whole length of the
abdomen, and dividing the black scales into two squarish areas on each segment ;
but it is commoner to find this longitudinal stripe distinguishable only on the last
two or three segments, and such specimens will sometimes have to be examined
carefully to avoid confusion with other species, such as O. punctor and O. communis.
The pro-epimeral scales will separate it immediately from all other Palaearctic
species.

An interesting variation is shown in a female from Szeged ( Kertész), which has the
abdomen almost entirely covered with yellowish scales, the black ones occurring only
towards the middle of some of the segments without forming definite markings ;
the mesonotal scales are also much paler than usual, especially towards the sides.
The pro-epimeral scales are normal for the species. A similar specimen from Budapest
has been described by Theobald (Mon. Cul. iv, p. 344) as Culex lutescens, but, as
stated above, I prefer to use Fabricius’ name in another sense.

Walker (List Dipt. Brit. Mus. i, p. 8) queries C. musicus as the same as
C. quadratimaculatus. The synonymy indicated may be correct, but the specimens
are no longer in existence.

The larva is one of the few in:this genus which lives through the winter, though
perhaps many of the eggs do not hatch until the early spring. There is some
individual variation in the number of hairs on the antero-dorsal side of the siphon.

Distribution.—West, central and south Europe. Locally common in England,
France, Belgium, Germany and Denmark. Also Italy (Rossi ; Taranto, Hargreaves ;
Livorno, Mann) ; Macedonia (Waterston). Not yet recorded from Sweden, Finland,
Russia or Asia.

13. Aédes (Ochlerotatus) lepidonotus, Edw. (figs. 5b, 5f, 8e.)
Ochlerotatus lepidonotus, Edwards, Bull. Ent. Res. x, p. 132 (1920).
Distinguished by the characters given in the keys and by the uniformly pale-
scaled abdomen of the female. The larva is unknown.
Distribution.—Macedonia (Waterston). f

14. Aédes (Ochlerotatus) albescens, sp. n.

Apparently related to A. lepidonotus, Edw., but the postnotum bears no scales.
Almost all the scales of the body whitish, except on the proboscis and palpi, where
they are mostly brown. Pro-epimeral scales whitish, rather narrow, some of them

MOSQUITOS OF THE PALAEARCTIC REGION. 309

curved. Palpi rather long, nearly one-third as long as the proboscis. Wings rather
scantily scaled (not rubbed); veins all pale, membrane whitish ; costal scales all
whitish-yellow ; first longitudinal vein with some dark scales. Integument of legs
pale yellow, scales almost all whitish-yellow, those on the terminal tarsal segments
browner ; no sign of pale tarsal rings. Claws all toothed. Integument of thorax

and abdomen blackish.
W. Srper1A: Omsk (Grand); 19. Type in Helsingfors Museum.

15. Aédes (Ochlerotatus) alpinus (Linn.).

Culex alpinus, Linnaeus, Flora Lapp. Ed. 2, p. 381 (1792).

Culex nigripes, Zetterstedt, Ins. Lapp. p. 807 (1838) ; Henriksen and Lundbeck,
Med. Groenland, xxii, p. 595 (1917).

Aédes innuitus, Dyar and Knab, Insecutor Inscitiae, v, p. 166 (1917).

Aédes (Ochlerotatus) nearcticus, Dyar, Rept. Canad. Arctic Exp. iui, Pt. C, p. 32
(1919),

Aédes alpinus, Dyar, Insecutor Inscitiae, viii, p. 52 (1920).

This species is fairly readily distinguishable by the unusually dense bristles on
the thorax, looking, as Dyar has remarked, “as though it had a long woolly coat
to keep out the cold.” The very dark colour of the mesonotal scales, the black
integument of the whole body, and the straight, shining greyish-white abdominal
bands are also characteristic. However, since the density of the bristles is some-
what variable (smaller specimens being less bristly), and they are also rather lable
to denudation (though less so than the scales), it is not always easy to distinguish
the species from A. cataphylla and A. parvulus. In fact A. alpinus might almost
be regarded as a race of A. cataphylla which has become adapted to arctic conditions
by the multiplication of its bristles and the thickening of the chitin of the
whole body.

I have failed to obtain a European male for examination from any correspondent,
and it is very unfortunate that the good series of the species brought from north
Russia by Capt. Carment and Dr. E. A. Cockayne consisted of females only. I
am, however, indebted to Prof. Sjéstedt and Dr. Lundbeck for several males from
south-west Greenland. These differ from the females (from the same place) in
showing hardly any scales on the thorax ; consequently they appear a good deal
blacker ; they are also even more hairy, especially on the abdomen. The hypopygium
very much resembles that of A. cataphylla, except in being far more heavily chitinised,
especially in the anal and genital parts. The apical lobe of the side-piece is very
small, hardly distinguishable ; the basal lobe has a moderately stout spine.

It seemed highly probable that these Greenland specimens were 4. innuitus,
D. & K., but that species was described,as having a “ double angular membrane ”
on the appendage of the claspette, as in A. lazarensis, while the males I have examined
showed the normal single membrane. Dr. Dyar therefore re-examined the types
of A. innuitus, and reports that the original description was in error ; the membrane
is really single, and the structure of the hypopygium of A. tmnutus and A. nearcticus
is really identical. That being so, there can be little doubt that 4. alpinus is also
the same, especially since we now know that a number of northern species of A édes
are common to the Old and New Worlds.

Distribution.—Probably a circumpolar species, occurring in all the lands to
the north of the Arctic Circle. I have seen males and females from south-west
Greenland, and females from Finmark, Lappland, Murmansk, and north-west
Siberia ; also a female from the Paris Museum labelled “ Fontainebleau, Dufour et
Laboulbéne.”” In regard to this last there must surely have been some mistake in
labelling.

310 F. W. EDWARDS.

16. Aédes (Ochlerotatus) detritus (Hal.) (fig. 7h).

Culex detritus, Haliday, Entom. Mag. i, p. 151 (1833).
Culex salinus, Ficalbi, Bull. Soc. Ent. Ital. xxviii, p. 29 (1896).
Culex terriet, Theobald, Mon. Cul. iii, p. 193 (1903).

This species is subject to a good deal of variation. Normally the dark parts
of the abdomen are speckled over with light scales, but these are sometimes absent.
Normally also the male palpi are entirely dark-scaled, but specimens are not
infrequently met with in which the long joint bears a good many pale scales on its
apical half. When both these variations occur together the specimens are not easy
at first sight to distinguish from A. salinellus. Apart from the structure of the
male hypopygium, which is of course diagnostic, the best distinctions are to be
found in the almost uniform brown colour of the mesonotal scales of A. detritus,
and the row of black spots down the middle of the venter, which are usually
conspicuous in living specimens, though they are not so easily seen in the dry.

So far as known, the larva is confined to salt or brackish water, and, as in many
other such forms, the gills are greatly reduced in size.

Distribution.—European coasts, from Ireland and France to Denmark and
Macedonia ; also coasts of North Africa, Suez Canal and Palestine. Apparently
does not occur inland in eastern Europe, as do its frequent associates A. caspius
and A. dorsalis. I have, however, seen a single male specimen, with a hypopygium
identical with that of European 4A. detritus, from Kashgar, Chinese. Turkestan
(Racquette, in Stockholm Museum).

17. Aédes (Ochlerotatus) cataphylla, Dyar, var. rostochiensis, Martini (fig. 7 1).

Aédes cataphylla, Dyar, Insecutor Inscitiae, iv, p. 86 (1916).

Aédes prodotes, Dyar, Insecutor Inscitiae, v, p. 118 (1917).

Aédes rostochiensis, Martini, Uber Stechmiicken, p. 246, diagnosis only (1920),

and Sitzb. u. Abh. natf. Ges. Rostock vii, p. 204 (1920). ~

This species and A. salinellus are together distinguished by the pale-speckled
femora and the more or less conspicuous pale ring towards the end of the long joint
of the male palpi ; they differ from A. detritus in never having scattered pale scales
on the dark parts of the abdomen, though the eighth segment is generally almost
entirely pale. The two species A. cataphylla and A. salinellus are extremely alike ;
the most obvious difference in the adult is in the colour of the scales of the proboscis,
which in A. cataphylla are almost all black, in A. salinellus extensively pale about
the middle, but I am not sure how far this difference is constant. There is a slight
but fairly well defined difference in the hypopygia (indicated in the key), but the
most striking character of A. cataphylla is the presence of a couple of disconnected
spines at the end of the pecten of the larval siphon.

None of the old European names seem to be applicable to this species, but I am
satisfied that it is specifically identical with the western North American A. prodotes,
Dyar (which Dyar now recognises to be the same as his previously described
A. cataphylla). The male palpi have the same pale ring; the larva has the same two
detached spines on the pecten, and the hypopygia are identical, except for a slight
difference in the size of the apical lobes of the side-pieces. The American specimens,
however, are on the average smaller, and the mesonotal scaling is rather different
from the usual European type, being rather darker in the middle and with more
numerous white scales at the sides. Nevertheless the Norwegian specimens in
the British Museum have the sides of the mesonotum even whiter than the
American specimens I have examined.

Distribution.—I have seen male specimens from Denmark (Wesenberg- Lund) ;
Sweden (Smaland, Stockholm and Dalecarlia, Boheman); Norway (Smaalenene,

MOSQUITOS OF THE PALAEARCTIC REGION. Fo |

Prof. O. Collet); Finland (Jamsa, Bergroth ; Helsingfors; Hattula, L. v. Essen ;
Finnstrom, Frey; Sund, Forsius ; Karislojo, J. Sahlberg; Kuusamo, Frey) ;
Germany (Frankfurt-Oder, Riedel; Berlin, Oldenberg); Austria (Prater, Mann:
this specimen was insufficiently examined and may have been A. salinellus) ; Siberia
(Yeniseisk, 58° 20’, Trybom).

18. Aédes (Ochlerotatus) salinellus, Edw. (fig. 7 k).

Aédes salinellus, Edwards (?) in Wesenberg-Lund, K. Danske Vid. Selsk. Skr.
Nat. Math. Afd. vii, p. 197 (1921).

Ochlerotatus salinellus, Edwards, Ent. Tidskr: ps’ 52: (1921).

Aédes terriei, Martini (nec Theo.), Uber Stechmiicken, p. 112 (1920).

The distinguishing points of this species have been mentioned under A. cataphylla.
In spite of their close similarity, there can be little doubt that the two species are
distinct. I am not satisfied that any older name can be applied to this species. The
type of C. fterriei is certainly nothing but a normal, if rather small, female of
_ A. detritus, which is common in the locality where C. terrier was taken. It is quite
possible that A. salinellus may eventually be found to be the same as some known
American species, but at present I have not been able to identify it with any such ;
it may be the European representative of A. impiger (=decticus, H.D.K.), which
has the male palpi all black and fewer bristles on the ninth tergite.

The name salinellus was suggested partly because of the strong resemblance to
A. detritus (salinus), and partly because the first specimens sent me by Dr. Wesenberg-
Lund were reared from brackish water.

Distribution.—Northern and central Europe. I have seen males from England
(Nottinghamshire, Carr) ; Denmark (Wesenberg- Lund) ; Germany (Berlin, Oldenberg,
Lichtwardt ; Kiel, Wiedemann); Austria (Vienna, Pokorny) ; Hungary (Budapest,
Kertész; Bethlen, Ujhelyi; Péstyén, Lichtwardt). Also some small females,
probably of this species, from south Russia (Waloniki, Velitschkousky). Martini records
it from various places in north Germany, generally near the sea, sometimes in
company with A. detritus.

19. Aédes (Ochlerotatus) diantaeus, H.D.Kk.

Aédes diantaeus, Howard, Dyar and Knab, Mosq. N. & C. Amer. iv, p. 758 (1917).
Aédes serus, Martini, Uber Stechmiicken, p. 96 (1920).

An extremely distinct species in larva and male hypopygium ; less well defined
in the female, but recognisable by the deep bluish-black colour of the tibiae and tarsi,
and the broad black stripe down the middle of the mesonotum. Owing to the
colour of the legs, and the usually interrupted pale bands of the abdomen, the female
might easily be mistaken for A. geniculatus, which of course differs in the structure
of the tip of the abdomen, as well as in thoracic markings. I have compared adults
and larvae‘of European and North American specimens, and can find no difference,
except in the length of the anal gills of the larva, which is known to be a variable
character. Neither of the published figures of the hypopygium is very accurate ;
in particular it should be noted that Martini’s omits altogether the conspicuous
hair-tuft of the side-piece.

Distribution —1 have examined the following European material :—Finland
(Kuust6, Kuusamo and Kiiminki, R. Frey) ; Denmark (Wesenberg- Lund). Martini’s
material presumably came from the Hamburg district.

20. Aédes (Ochlerotatus) sticticus (Meigen).

(2) Culex lateralis, Meigen, Syst. Beschr. i, p. 5 (1818).
Culex sticticus, Meigen, Syst. Beschr. vii, p. 1 (1838).

SZ — F. W. EDWARDS.

Culex concinnus, Stephens, Illustr. Brit. Ent. Suppl. p. 19 (1846).

Culex nigripes var. sylvae, Theobald, Mon. Cul. 11, p. 96 (1901).

Culicada sylvae, Theobald, Mon. Cul. iii, p. 194 (1903).

(?) Aédes aldricht, Dyar & Knab, Proc. U.S. Nat. Mus. xxxv, p. 57 (1908).

Culicada nigrina, Eckstein, Centralbl. f. Bakt. Abt. Orig. Ixxxii, p. 67 (1918).

Culex nemorosus var. dorsovittatus, Villeneuve, Bull. Soc. Ent. France, p. 57
(1919).

Ochlerotatus dorsovittatus, Séguy, Bull. Mus. Paris xxvi, p. 408 (1920).

(?) Ochlerotatus lesnet, Séguy, Bull. Mus. Paris xxvi, p. 328 (1920).

Culicada nemorosa salina, Brolemann (nec Ficalbi), Ann. Soc. Ent. France,
Ixxxviil, p. 81 (1919).

This species is fairly easily recognisable by the general blackish colour, the
white scales on the sides of the mesonotum, contrasting strongly with the dark
brown stripe which occupies the middle third, and the whitish stripe on the outer
side of the hind tibia of the female (often, though not always, present also in the
male). The hypopygium (figured by Séguy and Brolemann) is characterised by
the extremely short appendage to the claspette, which is very little longer than
broad, and not much broader than the width of the stem, and by the large basal
lobes, which are to a great extent separated from the side-pieces.

The species varies a good deal in size, though it is on the average smaller than
O. punctor, to which it is most nearly allied. It also varies in regard to the abdominal
bands of the female, which are in some specimens reduced to lateral spots (as in
the type of C. sticticus, and as described by Eckstein for C. nigrina), while in others
they are complete and scarcely even contracted in the middle (as in the types of
C. concinnus and C. sylvae). I do not believe that these differences represent anything
more than individual variation. Eckstein describes and gives rough figures of a
difference between the hypopygia of his C. mzgvina and what he regards as C. lateralis,
stating that the appendage of the claspette is shorter and less broad in the latter
than in the former. This may be true, and if so must indicate that we are dealing
with two distinct species, but I have seen no specimens corresponding to Eckstein’s
figure of C. lateralis, and am inclined to think he has merely shown the same structure
from two different points of view.

Since there may be two allied species here, and since there has been difference
of opinion as to the interpretation of Meigen’s C. lateralis, Theobald using the name
for A. gentculatus, it does not seem advisable to adopt this earlier name for the
species ; the type of C. lateralis being no longer in existence, it seems best to follow
Theobald in adopting Ficalbi’s suggestion that it is the same as C. albopunctatus,
Rond. (A. geniculatus). Of C. sticticus there is a female in good condition in Meigen’s
collection at Paris, and Séguy reports that the hind tibiae have a distinct pale stripe
on the outer side. The name sétcticus may therefore be used without further question.

I have examined the types of C. concinnus and C. sylvae and cotypes of
C. dorsovittatus, and have no doubt as to their identity. Séguy’s figure of the
hypopygium of.O. lesnei represents a structure apparently identical with that of
A. sticticus, his other figure of O. dorsovittatus being rather inaccurate, besides showing
the same structure in a different position. Specimens of A. séecticus were collected
by M. Lesne in the same locality and at the same time as the type of O. lesnet, and I
cannot help thinking that Séguy has confused two different species in his description:
the tarsi of O. lesnet are said to have pale rings.

The North American species A. hirsuteron (Theo.), A. aestivalis, Dyar, and
A. aldrichi, D. & K., are all very similar to A. sticticus in coloration, as well as in the
male hypopygium, all having the same peculiar structure of the basal lobes. It is
not at all unlikely that A. sticticus occurs in North America under one or other of
these names. I can see no difference between European A. sézcticus and American
A. aldrichi, and consider that they are most probably identical, though distinctions

MOSQUITOS OF THE PALAEARCTIC REGION. 313

may be discovered on closer study, especially when the European form is better
known.

According to Eckstein the larva of A. nigrinus lives in flooded meadows together
with A. vexans and A. dorsalis, and several generations are passed through during
the year. This does not accord with what is known of the habits of the allied
North American species.

Distribution—Widely spread in Europe, but seems to be nearly always rare.
I have seen specimens (mostly females) from Scotland (Stephens’ type ; also Aberfoyle,
Carter) ; England (New Forest, Theobald’s types); France (Melun, Debrewzl ;
Bois de Lutterbach, Bois de Borne, Chalampe, P. Lesne) ; Germany (Liiben, Rados,
Halle, Loew ; Bonn, Schneider ; Berlin, Lichtwardt) ; Denmark (Jutland, Wesenberg-
Lund); Austria (Linz, Kahlenberg, Aigen, Mik ; Dornbach, Handlirsch ; Prater,
Pokorny ; Styria, Mann); Hungary (Szévata, Cstki); Siberia (Asinovo, 61° 25’
Trybom).

A long series collected by Dr. K. Kertész at Fuzine and Jasenak (Croatia) shows
very little variation ; in all these the abdomen of both sexes has complete white
basal segmental bands, and the dark-scaled area in the middle of the mesonotum
is more extensive than usual, the specimens agreeing in this respect with many
American examples of A. aldricht, though not showing a pale median line dividing
the dark area.

21. Aédes (Ochlerotatus) punctor (Kirby) var. meigenanus, Dyar (fig. 7 g).

Culex punctor, Kirby, Fauna Boreali-Americana, Zool. Ins. p. 308 (1829).

Culicada (or Ochlerotatus) nemorosus, Theobald, de Meijere, Edwards, Lang
(nec Meigen).

(2?) Culicada nemorosa {. haplolineata and f. alineata, Schneider, Verh. Nat. Ver.

Bonn, Ixx, p. 37 (1913).

Aédes punctor, Dyar, Insecutor Inscitiae, viii, p. 3 (1920), and ix, p. 71 (1921).

Aédes meigenanus, Dyar, Insecutor Inscitiae, ix, p. 72 (1921).

Aédes sylvae, Martini (nec Theobald), Uber Stechmiicken, p. 108 (1920).

This species is a variable one, and hence difficult to distinguish with certainty
from its allies. The absence of a definite speckling of pale scales on the femora and .
tibiae, and the creamy tint of the abdominal bands—those on the last few segments
being rather conspicuously narrowed in the middle—are, taken together, the best means
of distinguishing the female. The male hypopygium is very distinct; the large,
prominent basal lobes and the short stem and strongly chitinised appendage of the
claspette suggest that the species is really more nearly related to the A. caspius
group than to the other members of the dark-legged group; this supposition is to
some extent borne out by the larval structure.

In the commonest type the mesonotum has brown or ochreous scales at the sides,
with a broad longitudinal dark brown band in the middle ; this form is fairly distinct
from other European species, though it might perhaps be confused with A. sticticus
or A. diantaeus (cf. the distinctions of those species). Frequently, however, the
dark central band of the mesonotum is either absent altogether or represented by

two narrow bands, and such specimens are difficult to distinguish from A. communis
or A. pullatus.

I have rejected the name nemorvosus for this species, because there appear to be
no examples of it so named in Meigen’s collection in Paris, while there is a male of
A. communis; the name nemorosus has been used to cover so many species that
there would be little advantage in retaining it. The present species is possibly
Meigen’s C. sylvaticus, but this is doubtful. While admitting that there are minute
differences in larva and adult, as well as some distinction in breeding habits, between
A. punctor and the European form, I cannot believe these are sufficient to justify

314 F. W. EDWARDS.

the maintenance of the two as distinct species. The European form should perhaps
be known as A. punctor var. meigenanus. The hypopygium is identical in the two
forms, and also in two or three other American forms which are regarded by Dyar
as distinct species. The relationship of all these forms appears to require closer
investigation.

It is quite possible that there may be more than one form in Europe, but the
only evidence I have seen of this is provided by a male from Kuusamo, Finland,
which has the hypopygium (and indeed the whole body) much more strongly chitinised
than usual, and the hairs on the apical lobe rather longer. This may perhaps
represent a» distinct variety or species.

Distribution.—Apparently widely spread throughout northern Europe, and
probably spreading across Siberia into Alaska and Canada. I have seen males from
3ritain; France; Belgium; Sweden (Stockholm, SmAaland, Boheman ; Oster-
edtland, Wahlberg) ; northern Lapland (Boheman) ; Vinland (Kuusamo, Finnstrém,
Frey ; Kuust6, Lundstrém ; Hattula, L.v. Essen ; Karislojo, J. Sahlberg ; Wiborg,
Pipping ; Jamsa, Bergroth ; Tvarminno, Levander) ; Germany (Berlin, Oldenberg,
Stobbe ; Brocken, Lichtwardl); Hungary (Munkacs, Ujhelyt); also females,
probably this species, from Austria (Hammern, M ik); Tyrol (Landro, Mann) ;
Siberia (R. Ob, Finsch ; Turuchansk, Yeniseisk, Inserovo, Antsiferovo and Nasimovo,
all on R. Yenisei, Tvybom).

22. Aédes (Ochlerotatus) parvulus, sp. n. (fig. 8 f).

3 Palpi all dark, long-haired, slightly shorter than the proboscis, proportions
of joints, 55: 26: 19. Head-scales mixed in the middle, then grey, then a patch
of blackish scales on each side ; the broad lateral scales extend further up towards
the middle of the head than usual ; bristles black. Thorax dark brown ; some white
scales at the sides of the mesonotum ; bristles rather dense. Abdomen black, the
tergites with narrow straight basal bands of pure white scales. Legs almost all
dark, the femora with only a few scattered pale scales and pale beneath. (Hind
legs missing.) Length 4-5 mm.

Hypopygium : Lobes of ninth tergite each with about six short hairs. Basal
lobe of side-piece prominent, rather pointed, with many long curved hairs but with-
out a spine. Apical lobe very small, with a few short straight hairs. Claspette
with the stem strongly curved; appendage with a broad membranous expansion
commencing near the base.
© Similar to the male. Palpi about one-sixth as long as the proboscis. The
hind tibiae have no pale lateral stripe. Length 3-5 mm.

A. parvulus has the appearance of a small, less hirsute race of A. alpinus, and
perhaps it really is so, but the less strongly chitinised hypopygium, the absence of a
spine on the basal lobes of the side-pieces, the slightly shorter male palpi, and the
less speckled femora seem to be sufficient to distinguish it specifically.

Distribution.—Finland: Kittila (¢ F. Silén, 2 U. Sahlberg); type male and
female in Helsingfors Museum; also several females, doubtfully conspecific, in
Helsingfors and British Museums, from Karislojo (J. Sahlberg), Kusomen. (Hellén)
and Suomussalmi (Hellén) ; these last have mostly a fairly distinct pale hind tibial
stripe, and the broad head-scales do not extend so far up as in the type.

93. Aédes (Ochlerotatus) communis (De Geer) (fig. 7 j).
Culex communis, De Geer, Mémoires, vi, p. 316 (1776).
Culex nemorosus, Meigen, Syst. Beschr. i, p. 4 (1818).
(2) Culex fasciatus, Meigen, Klass. i, p. 4 (1804).

(?) Culex leucomelas, Meigen, Klass. i, p. 3 (1804).

MOSQUITOS OF THE PALAEARCTIC REGION. 315

(?) Culex sylvaticus, Meigen, Syst. Beschr. i, p. 6 (1818).

Aédes obscurus, Meigen, Abbild. Zweifl. Ins. pl. ii, fig. 2 (1830).

Culex lazarensis, Felt & Young, Science, xx, p. 312 (1904).

Culicada nemorosa forma dtplolineata, Schneider Verh. Nat. Ver. Bonn, Ixx,
pees (1913).

(?) Aédes tahoensis, Dyar, Insecutor Inscitiae, iv, p. 82 (1916). -

(2?) Aédes pionips, Dyar, Insecutor Inscitiae, vii, p. 19 (1919).

Ochlerotatus palment, Edwards, Ent. Tidskr. p. 52 (1921).

This is one of a group of species which can only be satisfactorily distinguished
by the structure of the male hypopygium. When this organ is mounted and
examined under the high power of a binocular microscope, A. communis may be
readily distinguished from other species by the two ridges at the base of the appendage
of the claspette. These are best seen when the whole hypopygium is viewed from
above ; the small additional ridge is on the outer side of the appendage and at the
base only. The long, strongly arched stem of the claspette and the form of the basal
lobes seem to show that A. communis is more closely related to A. cataphylla and
A. salinellus than to A. pullatus or A. punctor, though in coloration the first two
species are more easily distinguished from A. communis on account of their speckled
femora and tibiae. The indications of relationship afforded by the male
hypopygium are also supported by the larval characters.

A. communis is evidently one of the most abundant woodland mosquitos of Europe,
and is no doubt the species which has most frequently been identified as Meigen’s
C. nemorosus. This, together with the fact that a male of the species is included
in Meigen’s series of C. nemorosus in the Paris Museum, will definitely settle this
name ; although Meigen’s description (brownish-yellow thorax, etc.) does not agree.
De Geer’s description of the adult and larva of C. communis, however, is quite sufh-
ciently detailed for identification, and I have therefore adopted his name for the
species. The disappearance of the name nemorosus from dipterological literature
will be an advantage rather than otherwise, since it is now known that a number of
species have been confused under this name.

I have examined the type of Meigen’s Aédes obscurus, which I received on loan
through the kindness of M. Séguy. The hypopygium is apparently identical with
that of A. communis, and the short palpi (if they were not merely broken) were
therefore probably an individual abnormality similar to those which I have recorded
as occurring in A. punctor.

In describing C. nemorosus, Meigen refers to his earlier description of C. veptans
(Klass. i, p. 3, a doubtful identification of Linnaeus’ C. reptans) as synonymous.
In this earlier description, however, the tarsi are said to be white-ringed, so that the
same species cannot have been referred to in both descriptions. The explanation
probably is that Meigen intended to give a reference to his C. leucomelas, very briefly
diagnosed immediately after C. reptans, with the remark, “ Diese Art, die vielleicht
nur eine Abandrung der vorigen ist, unterscheidet sich von derselben bloss durch
die ganz schwarzen Fiisse.” This name C. leucomelas has been overlooked, and
will unfortunately necessitate the renaming of a South American species.

Meigen’s C. sylvaticus (fasciatus, 1804) is impossible to determine from the
description, and I see no particular justification for Martini’s suggestion that it is
Culex apicalis; since the type does not exist it will be as well to accept Meigen’s
statement (Syst. Beschr. vi, p. 241) that it is only his C. memorosus.

The American forms A. lazarensis and A. tahoensis, and probably also A. pionips,
differ in such minute details that they can hardly be ranked as more than varieties
of A. communis, but it is interesting to note that the hypopygium of the Alaskan
tahoensis is the more nearly identical with European communis in regard to the exact
position of the spine on the basal lobe of the side-piece, the only point in which Dyar
has indicated distinctions between the American forms. I had intended to describe

316 F. W. EDWARDS.

a new species, A. palment, on account of some differences which I thought I perceived
in the male hypopygium in two specimens from Finland. On a re-examination I
failed to verify these differences, but meanwhile, unfortunately, I had published the
name palmeni in my key to the Swedish species.

Distribution.—Europe, except west and south, and probably across Siberia to
Alaska and Canada. I have examined male specimens from Denmark (Wesenberg-
Lund); Sweden (Dalecarlia, Vesterbotten, Norrbotten, Boheman); Finland
(Helsingfors, J. Sahlberg; Seitjaur, Palmen); France, Forét de Marles, Alluaud) ;
Germany (Berlin, Lichtwardt); Austria (Admont, Steiermark, Strobl ; Richenau
and Linz, Mzk ; Dornbach, Handlirsch).

I have also seen females, probably of this species, from Lappland and Siberia
(Yeniseisk).

24. Aédes (Ochlerotatus) pullatus (Coq.) var. jugorum (Villen.) (fig. 7 i).
Culex pullatus, Coquillett, Proc. Ent. Soc. Wash. vi, p. 168 (1904).
Culex jugorum, Villeneuve, Bull. Soc. Ent. France, p. 58 (1919); Séguy, Bull.
soc. Ent. France, p. 39) figs. (1921).
Aédes metalepticus, Dyar, Insecutor Inscitiae, viii, p. 51 (1920).
Aédes gallii, Martini, Uber Stechmiicken, p. 110 (1920).

This closely resembles A. communis, and it is almost impossible to distinguish
the two satisfactorily, apart from the male hypopygium and the larva, which are
very distinct. A. pullatus is somewhat smaller than A. communis, the integument
is perhaps darker, and there are more numerous white scales on the mesonotum.

I have not seen the types, but an Italian male of A. metalepticus sent by Prof.
Bezzi is practically identical in structure with an American male of A. pullatus in
the British Museum ; the distinctions given by Dyar must be due either to individual
variation or to differences of mounting. The only differences observable in the Italian
specimen were that the stem of the claspette was slightly stouter, slightly more
angulated, and with a more distinct bristle arising from the angle ; the main spine
of the basal lobe was somewhat stouter, and the pale scales of the thorax whiter.
These differences taken together may possibly indicate a varietal distinction.

Dyar suggested that C. gugorum might be the same as A. metalepticus, and this
is almost certainly the case, but Séguy’s figure of the hypopygium of a specimen
(not the type) of C. jugorwm shows a distinct apical hair-tuft on the side-piece, as
in A. intrudens. The Verestorony examples that I have examined have an aggrega-
tion of hairs in this position, almost suggesting a tuft, and Séguy may have exaggerated
the appearance of a tuft in his specimen, or the species may be somewhat variable.
Both Séguy’s and Kertész’s specimens agree with Bezzi’s except in this one point.
As remarked by Villeneuve, there are long dense hairs arching over the upper (sternal)
surface of the hypopygium, as in A. vusticus, A. cataphylla, and some others.

Distribution—As yet imperfectly known; occurs in mountainous regions of
Europe and North America, and therefore probably also of Central Asia. Recorded
by Martini from Switzerland (Galli-Valerio), by Dyar from north Italy (Bezz1), and
by Villeneuve from the Pyrenees (Byolemann). I have seen males from north Italy
(Scais, Bezzt); Transylvania (Verestorony, Kertész); and the Balkans (Vermosa,
1200m., Greuze, Penther); also females, possibly of this species, from north Sweden
(Kiruna, Lichtwardt).

25. Aédes (Ochlerotatus) intrudens, Dyar (fig. 8 f).
Aédes impiger, Howard, Dyar and Knab (nec Walker), Monogr. iv, p. 755 (1917).
Aédes intrudens, Dyar, Insecutor Inscitiae, vii, p. 23 (1919).
I know of no satisfactory means of distinguishing the adults of this species from
those of A. pullatus or A. communis, but the hypopygium is quite distinct, owing

MOSQUITOS OF THE PALAEARCTIC REGION. 317

to the dense tuft of hairs near the tip of the side-piece, projecting caudally from
about the level of the apical lobe, though not actually from the lobe. The long
hairs arching over the upper surface are not nearly so numerous as in A. pudllatus.
A. diantaeus also has a conspicuous hair-tuft, but in that species the tuft is situated
about the middle of the side-piece and projects inwards. The claspette-stem in
A. intrudens is angulated, with a projection arising from the angle, and a bristle
on the projection.

Distribution.—Widely spread in Canada. I have seen only one European male,
in the Berlin Museum, labelled 16.v.44, H. Loew. Dr. Enderlein informs me that
Loew was most probably in the Posen district on this date.

Subgenus Finlaya, Theo.

Adult. Proboscis slender, longer than the front femora. Palpi short in the
female ; from one-half to nine-tenths as long as the proboscis in the male, the last
two joints usually slightly thickened, moderately hairy and turned downwards.
Head scales and ornamentation of thorax variable. Lower mesepimeral bristles
absent. Male hypopygium with well developed claspettes, which bear a long and
rather slender appendage; no apical or basal lobes to side-pieces ; clasper and
aedoeagus as in Ochlerotatus. Eighth segment of female only partly retractile,
the sternite large and prominent in repose; cérci rather short. Front and middle
claws of female toothed, the hind pair simple.

Larva. As in Stegomyia, with some exceptions.

Most if not all the members of this subgenus breed in tree-holes and similar
situations or in rock pools. The species are most numerous in the Oriental region,
but they are found in practically all parts of the world, except the polar regions.
Owing no doubt to their restricted breeding-habits, there is a strong tendency in this
subgenus to the production of local species, and comparatively few have a wide
distribution.

Adults.

1. Tarsi (at least hind pair) with whitish rings .. fi, a se Say
Tarsi all dark .. st os ie #2 ‘1 si <i obi aoe

2. Tarsal rings extending on to apices of joints; thorax not distinctly lined
togot (Theo.).
Tarsal rings at bases of joints only ; thorax conspicuously lined .. a3 8

3. First three joints of hind tarsi ringed at base, last two all dark
japonicus (Theo.).
All joints of hind tarsi ringed at base ea He koreicus, Edw.

4. Head scales almost all broad and flat; a large (sometimes divided)
silvery-white area on the front of the mesonotum .. miveus (Ludlow).
Head scales narrow except at the sides ; mesonotum otherwise .. ee

5. Mesonotum with narrow lines of yellowish scales, sides not broadly
whitish .. 5 ae ss ae eatont (Edw.).
Sides of mesonotum broadly whitish scaled a is - LP eS
6. Scales of scutellum mostly or all narrow and ochreous ..geniculatus (Oliv.).
Scales of scutellum all broad, flat and white ag Nc echinus, Edw.

Male Hypopygia.

1. Stem of claspette extremely short, knob-like. . sts sia togot (Theo.).
Stem of claspette about as long as the appendage is 5s ee
2. Side-piece with a tuft of large scales on the upper side .. niveus (Ludlow).
Side-piece with small scales on the outer side only .. 3

3. Side-piece with very long and rather dense hair, as long as the side-piece
itself, on the upper flap 38 et ede uh eatont (Edw.).
Hair on side-piece shorter, only about half as long as the side-piece as 4
(4183) z

318 F. W. EDWARDS.

4. Lobes of ninth tergite with 6-8 fine hairs .. japonicus (Theo.) ; koreicus, Edw.
Lobes of ninth tergite with 24 stouter hairs es geniculatus (Oliv.).

Since most of the larvae are unknown, a table cannot be given.

26. Aédes (Finlaya) togoi (Theobald).
Culicelsa togot, Theobald, Mon. Cul. iv, p. 379 (1907).

This is very distinct from the other Palaearctic species of the subgenus, both
in the tarsal markings and in the male hypopygium, which shows the following
characters :—Side-pieces a little over twice as long as their depth at the base ; upper
flap much expanded basally, its inner margin with a dense row of long, slightly
flattened bristles or bristle-like scales, extending the whole length except on the
expanded basal portion ; the lower flap has an aggregation of hairs at the base, but
no definite basal lobe. Claspers moderately long, nearly cylindrical, tapering ;
terminal spine not very long. Claspettes with the stem extraordinarily short,
reduced to a mere knob; appendage sickle-shaped, slender, not expanded in the
middle. Lobes of ninth tergite small, with about 6-8 moderately long hairs.

The male palpi are about three-quarters as long as the proboscis, nearly straight
and slightly hairy.

The larva is undescribed (unless recently by Yamada); Dr. Lamborn found
them in granite basins in cemeteries.

Distribution Japan (Osaka, Theobald ; Tokio, S. Yamada, Harmand ; Yoko-
hama and Kobe, Lamborn) ; eastern Siberia (Vladivostok, per Dr. C. S. Ludlow).

27. Aédes (Finlaya) japonicus (Theobald).

Culex japonicus, Theobald, Mon. Cul. i, p. 385 (1901).

Apart from the tarsal characters, which are diagnostic, this species is noteworthy
for the fine yellowish lines on the mesonotum ; similar lines are found in A. koreicus
and A. eatoni. The male palpi are slightly shorter than the proboscis, slender,
the last two joints somewhat upturned, and with rather scanty hairs, thus approach-
ing very nearly to the type of palpi found in Stegomyia. The male hypopygium is
extremely similar to that of A. geniculatus. Theobald suggested that this might
possibly be the same as A. aureostriatus (Dol.), but this is unlikely.

The larva is undescribed, unless recently by Yamada; Dr. Lamborn found a
few in similar situations to those in which he took A. togot.

Distribution.—Japan (Tokio, Theobald, Yamada; Kofou, L. Drouard de Lezey ;
Chuzenji, E. Gallots ; Nagasaki, Lamborn).

28. Aédes (Finlaya) koreicus, Edw.
Ochlerotatus (Finlaya) koreicus, Edwards, Bull. Ent. Res. vii, p. 212 (1917).
This should probably be ranked as a variety of A. japonicus, from which it differs
in little else than in having narrow white rings at the bases of the last two hind tarsal
joints. The palpi of the type male are perhaps slightly less hairy than in A. japonicus.
The larva is unknown.
Distribution.—Korea (Dr. R. G. Mulls).

29. Aédes (Finlaya) niveus (Ludlow).

Stegomyia nivea, Ludlow in Theobald, Mon. Cul. iii, p. 139 (1903).
Stegomyia pseudonivea, Theobald, Ann. Mus. Nat. Hung. iii, p. 75 (1905).
Stegomyia albolateralis, Theobald, Rec. Ind. Mus. ii, p. 289 (1908).

In typical examples of A. niveus the silvery area on the scutum forms a solid
patch, but in some specimens (generally females) it is divided more or less com-
pletely by a dark median stripe, which may even reach the front margin. The

MOSQUITOS OF THE PALAEARCTIC REGION. 319

specimens with this dark stripe (such as the Japanese examples recorded below)
may represent a distinct species or variety (albolateralis, Theo.), but in view of
Leicester’s remarks on the variability of the species I doubt if this is so. Unfortunately,
I did not examine the hypopygium of the Japanese male. A male from the Andaman
Islands shows the following characters :—Side-pieces about twice as long as their
depth (or three times as long as their width) at the base, tapering, densely scaly
on their outer and under sides, finely hairy on the inner side, some long hairs at the
base beneath ; a dense row or tuft of very long and broad scales on the upper (sternal)
side, not reaching the base. Claspers short, with a very long terminal spine which
is quite two-thirds as long as the clasper itself. Lobes of ninth tergite with about
four very long bristles. Claspettes with the appendage moderately expanded in
the middle, about equalling the stem in length. The male palpi are of practically
the same length as the proboscis, the last two joints turned downwards and bearing
long hairs.

If Eichwald’s Culex niveus is an Aédes, the name is ineligible for this species,
and pseudoniveus should be used instead, but in view of the uncertainty there
is perhaps no necessity to make the change.

The larvae are not yet described, though Leicester records finding them in cut
bamboos.

Distribution—A _ widely-spread Oriental species extending into Japan (Tokio,
S. Yamada; 1 $192 in coll. M. Koitdzum1).

30. Aédes (Finlaya) eatoni (I-dw.).

Ochlerotatus eatont, Edwards, Bull. Ent. Res. vi, p. 358 (1916).

Apart from the very different thoracic markings, this species differs from A.
geniculatus in the coloration of the hind femora, which are black above for their
whole length, instead of entirely white on the basal half. The female and larva
remain unknown, the description being based on a single male, the palpi of which
are only about two-thirds as long as the proboscis.

Distribution. —Madeira Island (Eaton).

31. Aédes (Finlaya) geniculatus (Oliv.) (figs. 1 b, 5g, 10 a).

Culex geniculatus, Olivier, Encycl. Méth. Hist. Nat. Ins. vi, p. 134 (1791).

(?) Culex lateralis, Meigen, Syst. Beschr. i, p. 5 (1818) et auct.

Culex ornatus, Meigen, Syst. Beschr. 1, p. 5 (1818).

Culex guttatus, Curtis, Brit. Ent. p. 537 (1834).

Culex fusculus, Zetterstedt, Dipt. Scand. ix, p. 3459 (1850).

Culex albopunctatus, Rondani, Bull. Soc. Ent. Ital. iv, p. 31 (1872).

The shining white, almost silvery knee-spots, and the coloration of the hind
femora will distinguish even somewhat rubbed adults of this species from any
member of the subgenus Ochlerotatus. Fresh specimens are obviously distinct by
their thoracic markings. From the allied A. echinus this species differs mainly
in the narrower scutellar scales; but even this is not absolutely diagnostic, as I
have seen a male from the south of France which is certainly only A. geniculatus,
but which has some small flat scales on the scutellum. The larva, however, is very
different from that of A. echinus, and as it is the only other species of the genus
which is found in tree-holes in Europe, no confusion with any other species is likely.

As I have previously stated (Ent. Mo. Mag. 1912, p. 277), I do not think there
can be any doubt that Olivier’s name should apply to this species, and that Theobald
was in error in using it for Culex hortensis. I now believe that the female which
Meigen originally described as C. ornatus must have been this species, although
the male which he described at a later date must have been something else, as
he refers to the white marks on the palpi. Van der Wulp, Verrall, Galli-Valerio

(4183) z2

320 F. W. EDWARDS. °

and others who identified C. ornatus as this species were therefore probably right.
Meigen’s C. lateralis may also be the same, as supposed by Theobald, but the
type being lost it is impossible now to say whether it was this species or A. sticticus.
Mr. J. E. Collin has kindly examined for me the two males of C. fwsculus in Zetter-
stedt’s collection at Lund, and from his notes and accompanying sketch of the claspette
there can be no doubt that C. fusculus is A. geniculatus. Zetterstedt must also
have included with these males rubbed females of other species, as one which was
sent me by Dr. Bengtsson in 1912 was an Ochlerotatus near O. cataphylla.

ee] a

yee ec

Fig. 10. First abdominal segment of larva of (a) Aédes geniculatus and (b) Aédes echinus, showing
the remarkable difference in the development of the hairs.

Distribution —Throughout Europe from France to Galicia and from south Sweden
to Macedonia, wherever there are deciduous trees, such as beech, sycamore, plane,
horse-chestnut, sweet chestnut, etc., in sufficient numbers. It occurs also in Corsica
(Yerbury, Séguy) and in Asia Minor (Brussa, Mann). It is not yet known from
Siberia or North Africa, though its known range will no doubt be extended by future
observations. The most northerly record I have is Scania, Sweden (Boheman).
Its absence from the far north may be accounted for by the fact that coniferous
trees, also birches and willows, do not readily form rot-holes that will contain water.
Its apparent absence from Scotland and Ireland is not so easy to explain. The
North American representative (A. tviseriatus, Say) is specifically distinct.

32. Aédes (Finlaya) echinus (Edw.) (figs. 6a, 10 b, 11).
Ochlerotatus (Finlaya) echinus, Edwards, Bull. Ent. Res. x, p 133 (1920).

The adult is very similar to A. geniculatus, apart from the two points mentioned
in the key, but the larva is strikingly different, chiefly on account of the remarkable
development of the hair-tufts on the thorax and abdomen. | These appear to corre-
spond rather closely in number and position with those of A. geniculatus, but the
component hairs are more numerous, distinctly plumose, somewhat longer, and
much stouter, giving the insect a very urchin-like appearance (whence the specific

MOSQUITOS OF THE PALAEARCTIC REGION. 321

name). Fig. 10 illustrates well the difference in vestiture of the first abdominal
segment in the two species; the following segments in both have the hairs longer.
(In the case of A. echinus both the skins preserved by Capt. Waterston are much
distorted, and careful reconstruction was necessary.) The larva of A. echinus
also differs from that of A. geniculatus in the distinctly longer antennae, which are
much more extensively pale towards the tip, in the much more numerous pecten
teeth, which are in a straighter row, in the somewhat smaller siphon and somewhat
larger comb teeth, as well as in some other details, perhaps not constant.

TERZI —

Fig. 11. Aédes echinus, Edw., end of abdomen of larva; comb and pecten teeth and mentum
more highly magnified.

The pupa differs from that of A. geniculatus in having the paddles more strongly
emarginate and the terminal hair placed at some distance outside the tip of the
midrib.

Distribution.—Macedonia (Stavros, Waterston ; reared from larvae in hole in
plane tree ; many adults of A. geniculatus also found in the same locality) ; Morocco
(Fez, Fowler); Algeria (Sergent).

OVA F. W. EDWARDS.

Subgenus Ecculex, Felt.

Adult.—As in Ochlerotatus, but the male palpi are usually no longer than the
proboscis; the vertex and the scutellum in many species are covered with flat
scales ; lower mesepimeral hairs are usually absent ; the male hypopygium has no
distinct claspettes, these being represented by hairy basal lobes ; while, on the other
hand, the claspers are highly modified in most species, and even in the simple forms
are distinguished by having the spine inserted before the tip. The aedoeagus is
quite different from that of Ochlerotatus, but almost identical with that of Aédes.
The hind claws are usually simple.

Larva.—Practically as in Ochlerotatus, but the pecten usually has detached teeth
outwardly, which in Ochlerotatus is rarely the case, and the siphonal tuft is usually
distinctly beyond the middle.

In E. vexans the frontal hairs are situated one in front of the other, as in Ochlero-
tatus, but in the rather numerous Ethiopian species of which the larvae are known
the anterior pair is placed outside the posterior, as in Culex and Aédes (s. str.).

It is only with reluctance, in deference to the opinion of Dyar, that I recognise
this as a distinct subgenus, as it is almost impossible to define apart from the characters
of the male hypopygium. However, there is no doubt that it is a natural group,
representing a distinct line of evolution in which the clasper has undergone
specialisation instead of the claspette. The separation 1s confirmed by present-day
distribution, the species being numerous in the Oriental and Ethiopian regions, only
a single one extending into the Palaearctic and Nearctic, while none are found in
the Neotropical, and only one or two in north Australia.

33. Aédes (Ecculex) vexans (Mzg.).

Culex vexans, Meigen, Syst. Beschr. vi, p. 241 (1830).

(2) Culex parvus, Macquart, Suites 4 Buffon, i, p. 36 (1834).

Culex articulatus, Rondani, Bull. Soc. Ent. Ital. iv, p. 30 (1872).

Culex malariae, Grassi, Atti Acc. Lincei, vii, p. 168 (1898) ; Noé, Bulle Soe) Ent:
Ital. xxxi, p: 244 (1899);

(?) Culex avabiensis, Patton (adult, not larva), J. Bombay Nat. Hist. Soc. xvi,
p. 633 (1905).

Apart from the characters of the male hypopygium and the somewhat shorter
male palpi, this species is not easy to distinguish from the annulipes group of the
subgenus Ochlerotatus, especially small specimens of A. maculatus. The structural
and scale characters are the same, except that the hind claws are often (not always)
simple ; in both the lower mesepimeral bristles are absent. We have therefore, in
the case of the female, to rely mainly on coloration for the determination of this species,
the most constant feature being the median emargination of the pale abdominal
bands. The white tarsal rings are always narrow, but rather variable ; in large
specimens they are sometimes not much narrower than those of the narrowest-banded
specimens of A. maculatus ; while in the other examples (generally small ones)
they are often so narrow as to be visible only under a lens. The head markings
- (the uppermost of the flat scales at the sides of the head being black, the rest pale)
are of a type very rarely met with in Ochlerotatus, but common in Ecculex, Stegomyta
and Finlaya. E. vexans differs from the great majority of species of the subgenus
in having no flat scales either on the vertex or on the scutellum, its resemblance to
the subgenus Ochlerotatus being thus increased.

Two varieties occur within the Palaearctic region: the typical form, in which
the abdomen has only the emarginate white bands on a dark ground; and the
variety nipponii, Theobald, in which the abdominal segments have, in addition to
the bands, a median whitish patch.

MOSQUITOS OF THE PALAEARCTIC REGION. 320

The species is so abundant in many parts of Europe as to constitute a serious
plague ; so much so that, according to Eckstein and others, agricultural work in some
districts has to be carried on by night and cattle stalled during the day, owing to the
attacks which A. vexans makes by day on the men and cattle.

The larva occurs characteristically in flooded meadows, and several generations
are passed through in the year. There is very little difference between the larva
and those of some species of Ochlerotatus which have detached teeth at the end of
the pecten, even the distinctions given in the key being somewhat doubtful. Accord-
ing to the figure and description in Howard, Dyar and Knab’s monograph (where
the species is named A. sylvestris, Theo.), the siphonal tuft is situated in the middle,
though according to Schneider and Martini it is placed well beyond the middle.
The only larvae I have seen are from Ceylon, and these agree entirely with
Martini’s description ; I suspect therefore that the American description may be in
error, as there is certainly no difference in the adults.

Distribution.—Apart from the domestic species (Culex pipiens, C. fatigans and
Aédes argenteus), this is the most widely spread ot all mosquitos, occurring practically
throughout the Palaearctic, Oriental and Nearctic regions. Possibly it may have
had its origin in tropical Africa, where there are a number of related forms, but if
so its apparent absence from that region at the present time is remarkable. It is
common throughout central Europe, perhaps less so in the south, and certainly rare
in the north. Some fresh records are: Sweden (Oeland I., Boheman) ; Finland
(Helsingfors, Frey ; Tvarminno, Levander) ; Italy (Susa, Sondrio, Torino, Macerata,
Chivasso, Bezzi); Asia Minor (Konia, Naday) ; Transcaspia (Tashkent, Aschabad,
C. Ahnger); Ussuri (Spasskaja, Wuorentaus) ; Persia (Enzeli, Buxton); Korea
(Yamada) ; Aden (Kazan Chand, per Capt. P. J. Barraud).

The variety mipponit occurs in China and Japan, also the Amur region (Ussuri,
Spasskaja, Wauorentaus).

Subgenus Aédes, Mg.

Adult. Proboscis (in the Palaearctic species) about equal in length to the front
femora, or slightly shorter. Palpi very short in both sexes. Antennae of the male
with the hair-whorls evenly spread all round the joints. Vertex with broad flat
scales, leaving only a small patch of narrow ones on the nape. Lower mesepimeral
bristles absent. Male hypopygium with the claspers deeply bifid, without terminal
claw, inserted before the tip of the side-piece ; the latter with small hairy basal lobes.
Aedoeagus with the parameres indistinct, almost membranous, the mesosome
chitinised in two lateral halves, which are split into rather numerous small spines ;
one very much larger spine is apically directed. Female cerci moderately elongate ;
eighth segment rather large. Front and middle claws of female toothed.

Larva. Antennae rather long, with numerous spinules and well-developed
tuft. Frontal hairs not one in front of the other, as in Ochlerotatus, but almost side
by side (in Lang’s terminology, the three post-antennal hairs are almost in one line,
the middle one not displaced). The median anterior thoracic tufts are absent.

34. Aédes (Aédes) cinereus, Mg.

Aédes cinereus, Meigen, Syst. Beschr. i, p. 13 (1818).

Aédes rufus, Gimmerthal, Bull. Soc. Imp. Nat. Moscou, xviii, p. 295 (1845).

Aédes leucopygus, Eysell, Abh. Ver. Naturk. Kassel, xlviii, p. 285 (1903).

Culex nigritulus, Zetterstedt, Dipt. Scand. ix, p. 3459 (1850).

Aédes fuscus, Osten-Sacken, Bull. U.S. Geol. Surv. iii, p. 191 (1877).

? Culex ciliaris, Linnaeus, Syst. Nat. Ed. xii, i, p. 1002 (1767).

This species need not be confused with any other in the Palaearctic fauna. The
mostly flat-scaled head, extremely short palpi of the male, reddish, unmarked thorax,

324 F. W. EDWARDS.

and dark-scaled dorsum of the abdomen of the female, should make it quite
unmistakeable. So far as European specimens are-concerned there is little variation,
though it should be noted that the thorax of the male is always much darker than
that of the female, usually quite black.

The larva does not differ in any very striking manner from those of the sub-
genera Ochlerotatus and Ecculex ; the main points have already been noted. The
early stages are spent usually in flooded meadows and large marshes, but the species
is also found in woods. It is commonly associated with A. vexans.

Distribution —Throughout Europe, and extending across Siberia to North America,
where it has a wide distribution. The following are some new records :—Italy
(Sondrio, Bezzi); Finland (various localities and collectors) ; Siberia (Yeniseisk,
58° 20’, and Turuchansk, 65° 55’, Trybom ; Omsk, Grano).

Subgenus Stegomyia, Theo.

Adult. Proboscis moderately slender, but stouter than in Ochlerotatus, scarcely
as long as the rather short front femora. Palpi short in the female, normally longer
than the proboscis in the male, the last two joints slender, upturned, with very few
hairs. Vertex with broad flat scales, few or no narrow ones on the nape. Thorax
usually with conspicuous and well-defined ornamentation. Lower mesepimeral
bristles absent. Male hypopygium usually without claspettes, unless these are
represented by hairy basal lobes; no apical lobes ; clasper with distinct terminal
spine. Aedoeagus divided into two more or less brush-like halves. Eighth segment
of female abdomen rather large, but distinctly retractile, the sternite not very
prominent in repose; cerci rather short. Front and middle claws of the female
either toothed or not.

Larva. Antennae short, with single hair and without spicules on shaft. Frontal
hairs single. Abdomen with or without numerous stellate tufts on dorsal surface ;
the eighth segment with a definite comb of teeth set in a single row. Siphon not much
more than twice as long as broad ; hair-tuft well developed and situated about the
middle.

The larvae of many African species live in tree-holes, leaf-axils, etc., and these
species show a much greater development of the abdominal hair-tufts than is seen
in the Palaearctic species.

Adults.
1. Mesonotum with a median silvery-white line He i Be In
Mesonotum without such line - ae me ote oe eS
2. Female claws simple (Japan) .. im ae a albopictus (Skuse).

Female claws toothed (Crete) .. re oh a cretinus, Sp. N.
3. Mesonotum with a lyre-shaped silvery-white mark ; tibiae dark except at
tip sit a a i oF ae ae argenteus (Poiret).
Mesonotum with four distinct white dots ; tibiae ringed with white a little
beyond the middle ; , vittatus (Bigot).

Male Hypopygia.
1. Clasper modified, swollen and hairy apically, the spine long and curved and
placed far before the tip ~~. - - ate vittatus (Bigot
Clasper normal ; spine shorter, straight, and terminal a Bs ar
2. Clasper shorter, narrowed at the tip, side-piece with a large, densely bristly
cred. Wipers se td 4 a avs os . argenteus (Poiret).
Clasper longer, slightly swollen at the tip, side-piece with a large, hairy
basal lobe af i vi ae by albopictus (Skuse).

).
2,

MOSQUITOS OF THE PALAEARCTIC REGION. 325

35. Aédes (Stegomyia) albopictus (Skuse).
Culex albopictus, Skuse, Ind. Mus. Notes iii, p. 20 (1895).
Stegomyia scutellaris, Theobald (nec Walker), Mon. Cul. i, p. 298 (1901).

This is the only species of Aédes in the Palaearctic region in which the front and
middle claws of the female are not toothed; it cannot however be removed from
the genus, or even from the subgenus Stegomyia, with which it agrees in all other
respects. The silvery line down the middle of the mesonotum will at once differentiate
it from all other mosquitos in the region except A. cretimus, its Mediterranean
representative.

The larva has been described and figured by Banks (Phil. J. Sci. A, iii, 1908,
p. 246) but he omits to notice the structural difference from A. argenteus in the shape
of the comb-teeth.

Distribution —A common semi-domestic species throughout the Oriental region,
occurring in Japan in the neighbourhood of Tokio (Yamada) ; Mt. Takao, near Hachioji
and Kofou (Paris Museum) ; Yokohama and Kobe, also Shanghai (Lamborn). It
occurs also in Madagascar and Réunion.

36. Aédes (Stegomyia) cretinus, sp. n.

Closely allied to A. albopictus, Skuse, but differs as follows :—A pair of small
round spots of white scales in the middle of the mesonotum, a little in front of the
wing-roots level with the posterior end of the central white stripe. Abdomen with
very distinct white basal bands on segments 2~7, somewhat narrowed in the middle.
Fourth hind tarsal joint darkened only at the extreme tip. Front and middle claws
toothed. As in A. albopictus, the front and middle femora have a narrow line of
white scales towards the base anteriorly, but no median white spot.

A single female in Herr Lichtwardt’s collection, labelled “ Creta. v. O. Culex
calopus, Mg.” Since, apart from the toothed claws, there are slight differences from
both A. albopictus, Skuse, of the Oriental region, and A. unilineatus, Theo., of Africa
and the Punjab, it is more likely that we are dealing with a distinct Mediterranean
representative of A. albopictus than that there has been any error in labelling.

Asecond female is in the Buda-Pest Museum from Amari, Crete, 4. vi. 1906 (Biro) ;
the abdomen and claws agree with the type, but the mesonotum is rubbed and the
hind tarsi missing.

37. Aédes (Stegomyia) argenteus (Poiret) (Sfegomyia fasciata).
Culex argenteus, Poiret, Journ. de Phys. xxx, p. 245 (1787).
Culex fasciatus, Fabricius, Syst. Antl. p. 36 (1805).
Culex calopus, Meigen, Syst. Beschr. i, p. 3 (1818).
Culex konoupi, Brullé, Exp. Sci. de Morée, Zool. ii, p. 289 (1836).
(2) Culex niveus, Eichwald, Reise Casp. Kauk. 11, p. 183 (1837).
Culex elegans, Ficalbi, Bull. Soc. Ent. Ital. xxi, p. 95 (1889).
Culex albopalposus, Becker, Mitt. Zool. Mus. Berlin, iv, p. 80 (1908).
Culex angustealatus, Becker, Mitt. Zool. Mus. Berlin, iv, p. 79 (1908).
? Culex aegypti, Linnaeus, Hasselquist’s Reise nach Palestina, p. 470 (1762).

The yellow fever mosquito is widely spread, though apparently nowhere very
abundant, in the warmer parts of the Palaearctic region. It occurs on the Atlantic
- islands, in Portugal, and all round the Mediterranean coasts. In the eastern
Mediterranean it is by no means confined to the coasts, since Barraud has found it
to be common at Aleppo. Further east it is known from Mesopotamia, Persia, and
Japan.

Trichwald says of Culex mniveus, which he records from Tiflis and Baku, ‘in
unzahliger Menge abends in den Zimmern bemerkt werden.’’ This, together with
his ‘“ thorax nigro alboque varius, alba pube obsitus’’ and “ pedibus nigro canoque
variis,” seems to suggest Aédes argenteus, but other parts of the description(“ alis

326 F. W. EDWARDS.

niveo-albis, corpore ex dimidio fere brevioribus,’’ and “ pedibus anticis in apice
utrinque fasciculo pilorum ornatis’’) will not apply to this or any other known
mosquito.

It is quite possible that Dyar may be right in identifying C. aegypti with this species.
Certainly it seems to be some Stegomyia, and A. argenteus is the only member of the
genus now known to occur in Egypt, but there are one or two points in the description
which quite definitely do not agree; I have therefore not adopted the name.

38. Aédes (Stegomyia) vittatus (Bigot).
Culex vittatus, Bigot, Ann. Soc. Ent. France, (4) i, p. 327 (1861).
Stegomyia sugens, Theobald, Mon. Cul. 1, p. 300 (1901).
Culex sugens, Wiedemann, Aussereurop. zweifl. Ins. 1, p. 545 (1828).

The white dots on the mesonotum, together with the white-ringed tibiae and
tarsi, make this species an extremely easy one to recognise. The pre-apical spine
of the male clasper, and the position of the siphonal tuft of the larva well beyond
the middle, suggest that the species may have more in common with the subgenus
Ecculex than with other species of Stegomyta, in spite of the spineless larval antennae
and the slender, bare, upturned male palpi. The species affords a good illustration
of the difficulty of drawing any hard and fast line between the subgenera of A édes,
and confirms the inclusion of all of them in one comprehensive genus.

The larva has been recorded as occurring in rock pools.

Distribution.—Corsica (Bigot). Also widely distributed in the Ethiopian and
Oriental regions, occurring as far south as Ceylon. It is remarkable that so
conspicuous a species has not been found in the Mediterranean region since
Bigot’s time.

Genus Armigeres, Theobald.

This genus is evidently closely allied to Aédes (especially the subgenera Aédes
and Stegomyia), so much so that scarcely any tangible differences can be discovered
in the adults. The proboscis is rather short (not longer than the front femora) and
is slightly but distinctly stouter throughout than in Aédes; also the tip is slightly
but distinctly curved downwards (at least in dry specimens), which is very seldom
the case in Aédes. In the allied Oriental genus or subgenus Levcesteria the
mesonotum is somewhat produced over the head, and this tendency is slightly
indicated also in the typical subgenus Avmigeres. The structure of the eggs and
manner of oviposition in Avmigeres is similar to that of Aédes; but Letcesteria
flava, according to Strickland, has peculiar egg-laying habits. The male clasper has
numerous spines, generally placed in a row (4-10 in Lezcesteria, 15-20 in Armigeres).
Another small point of distinction from Aédes is that the middle claws of the male
are apparently always equal and simple.

The main reason for keeping Aymigeres distinct from Aédes is the structure
of the larval siphon, which has only a minute and often scarcely distinguishable
hair-tuft, andno trace ofa pecten. This latter point constitutes such a sharp difference
from Aédes that the separation from that genus may be justified, in spite of the
feeble characterization of the adults. The anal gills are of large size and rounded
apically.

The genus is endemic in the Oriental region, a single species extending into Japan
and thus claiming our attention in this paper.

Armigeres obturbans (Walker).

Culex obturbans, Walker, Proc. Linn. Soc. London, iv, p. 91 (1860).
Culex subalbatus, Coquillett, Proc. U.S. Nat. Mus. xxi, p. 302 (1898).

MOSQUITOS OF THE PALAEARCTIC REGION. S27

Like the other species of the genus, this is a dark-coloured insect, with entirely
dark tarsi and mainly white venter, and with flat scales, mostly dark, covering the
head and scutellum. It differs from all its congeners in having a distinct if narrow
band of black scales at the apex of each abdominal sternite. The male palpi resemble
those of Stegomyia, but are entirely dark. I am indebted to Dr. H. G. Dyar for
information as to the identity of Coquillett’s type.

The larvae live in bamboo stems, and have been described by Banks (Phil. J.
Sci., A. ili, p. 240, 1908). They have remarkably large, sausage-shaped anal gills,
which enable them to remain long periods at the bottom.

Distribution.—Japan (Kofou, L. Drouard de Lezey ; Kouy-Tchéou, Fortunat ;
Hakone, E. Gallois ; Tokio, Yamada); also throughout the Oriental region, and
extending into Celebes, New Guinea and North Australia.

Genus Lutzia, Theobald.

Owing to the highly modified larval mouth-parts and antennae, and the peculiar
Siructure of the siphon and anal segment, it was long ago proposed by Christophers
to separate the Old World species of this genus from Culex as a distinct genus
( Jamesia) ; the same characters were used by Dyar and Knab in separating the
New World Lutzia from Culex. In revising the African CULICIDAE in 1912 I did
not accept this separation, owing to the apparent structural identity of the adults.
I now find, however, that an excellent diagnostic character exists in the numerous
lower mesepimeral bristles of Lutzia, and I therefore propose to revive this name.
There is no real difference between the Old World and New World forms, and I
consider Dyar’s separation of Jamesia and Lutzia on a small detail of aedoeagal
structure to be quite unjustifiable. The Old World species are all very similar,
their separation resting on small differences of colour and venation.

Lutzia vorax, sp. n. (fig. 5d).

Penultimate joint of male palpi with the integument and the hairs dark except
at the extreme tip. Abdominal tergites in both sexes all with rather narrow but
distinct apical pale ochreous bands. Lobes of mesosome of male aedoeagus enlarged
beneath a little beyond the middle, the enlargement with some minute teeth ; lobe
of side-piece with three strong spines only. The whole of the outer side of the hind
femora has the light and dark scales about evenly mixed. Cross-veins either in a
straight line, or else m-cu (posterior) placed beyond r—m (mid).

L. concolor (R.—D.), Theo., the commonest form in the Oriental region, differs
in having the last few abdominal segments entirely yellow-scaled, the yellow bands
on the anterior segments narrower; the lobes of the mesosome are not enlarged
beneath ; the lobe of the side-piece usually has a fourth spine more or less developed,
separate from the other three ; the outer side of the hind femora is entirely pale at
the base, from which a more or less definite pale line runs almost to the apex ; and
the cross-vein m-—cu is placed at least slightly before r—m.

L. halifaxi (Theo.), known from the Malayan region and Queensland, has the
hypopygium almost identical with that of L. vorax, the enlargement of the mesosomal
lobes perhaps more prominent and practically in the middle; it differs in having
the integument and hairs on the apical half or more of the penultimate joint of the
male palpi paler than the basal part, and in having few or no pale scales on the apices
of the abdominal tergites ; the hind femora are as in L. vorax, but darker ; the cross-
veins, on the other hand, are placed as in L. concolor.

Dr. Lamborn found the larvae in old cess-pits preying upon Culex fatigans.

Distribution.—Japan (Tokio, Yamada ; a series presented to the British Museum
in 1916, determined at the time as Culex concolor; the type of the new species is
one of the three males in this series; also Karuizawa, Cornford, and Nagasaki,

328 F. W. EDWARDS.

Lamborn). North India (Punjab, Barrow ; female only). Probably widely distri-
buted in the Oriental region, but confused with the two species above mentioned
and with the Ethiopian L. tigripes.

Genus Culex, L.

This genus, I find, is sharply distinguished from almost all other mosquitos by
the possession of distinct pulvilli. It is remarkable that the presence of these structures
has been overlooked for so long; Howard, Dyar and Knab even state positively
that they are absent throughout the family ; these and other writers must either
have omitted to study Culex closely, or else have used an insufficient magnification.
I have examined a large number of species of this genus, and find pulvilli present
in all ; they do not vary much in size, but are naturally more easily detected in the
larger species. Figs. 5d and 5e (made with the aid of a camera lucida) show clearly
the different appearance under a sufficiently high power between a hairy empodium
and a pair of true pulvilli. In the front and middle tarsi of the male the pulvilli,
like the claws, are elongated, and therefore less noticeable ; they may be seen,
however, on the hind tarsi as well as on all the feet of the female. The only other
mosquitos which possess pulvilli are the genera which on other grounds have already
been regarded as close allies of Culex: Culiciomyia, Lophoceratomyia, Micraédes
Carrollia, Lutzia, and Deinocerites (including Dinomimetes). The first three or four
of these should not be regarded as more than subgenera of Culex, though the last
two may be treated as distinct genera.

The following characters are also common to most if not all species of Culex ;
some of these will further help to distinguish the members of this genus from A édes :
Eyes very narrowly separated or even touching for a considerable length above the
antennae. Proboscis not or scarcely longer than the front femora. Male palpi
when long always slender, with the last two joints upturned. Male antennae always
plumose, with the hairs spreading out evenly all round. Spiracular and post-spiracular
bristles absent. Usually only one lower mesepimeral bristle or none; very rarely
two or three. Female abdomen blunt-ended, the cerci short and broad, eighth
segment not at all retractile. Male hypopygium without claspettes or basal lobes
to the side-pieces, but with subapical lobes bearing modified bristles. Tenth sternites
ending in a tuft or comb of spines. Mesosome a paired structure with pointed pro-
cesses. Claspers articulating in a more or less vertical plane. First joint of hind
tarsus as long as the tibia or slightly longer. Female claws always simple. Wings
with distinct microtrichia on the membrane ; cell R, markedly longer than its stalk
in the female ; vein A, ending much beyond the level of the base of R,.

Larva.—Antennae with a distinct hair-tuft, which is generally well beyond the
middle, the part of the antenna beyond the tuft usually rather suddenly narrowed,
and with few or no spinules; two long preapical spines. Hairs of mouth-brush
simple. Frontal hairs rarely if ever single, and never placed one in front of the other.
Anal segment with a complete chitinous ring (in the fourth stage only). Siphon
with numerous ventral tufts, or else greatly elongate.

The genus is essentially tropical and sub-tropical, only a very few species
extending into the temperate regions. Only C. apicalis and the domestic
C. pipiens and C. fatigans are common to Europe and North America.

Three fairly well-marked subgenera occur within the Palaearctic region, as
indicated in the following keys.

Adults.
1. First joint of hind tarsi distinctly shorter than the tibiae; small
obscurely coloured species (Barraudius) .. 3 Akane 34

First joint of hind tarsi scarcely, if at all, shorter than the tibiae wt BGS

14.

15.

16.

17)

18.

MOSQUITOS OF THE PALAEARCTIC REGION. 329

. Abdominal tergites with continuous lateral pale stripes .. modestus, Fic.

Abdominal tergites, with basal lateral pale patches. . <6 pusillus, Mcq.

. A row of small flat white scales round the margin of the eyes (Culicomyia)

impudicus, Fic.
Scales on the top of the head all narrow (Culex)... = 5s

. Prothoracic lobes and pro-epimera with numerous broad flat scales ; pale

bands of abdominal tergites apical (occasionally reduced to lateral spots) 5
Prothoracic lobes and pro-epimera with few or no flat scales sy 8

. Hind tibia with a distinct white spot on the outer side at the tip, hortensis, Fic.

No such spot... es oe 2 ei 5 .. apicalis, Adams.

. Dorsum of abdomen uniformly dark brown ; species without ornamentation

hayasu, Yam.

Abdominal tergites at least with basal lateral patches of pale scales a:
. Proboscis and tarsi pale-ringed. . ue be deg zk ,40 ae
Proboscis and tarsi without pale rings os Ai hs as .. 14

. Anterior two-thirds, or at least the middle third, of the mesonotum

with whitish scales, which contrast shar Ply with the dark scales of the
posterior third a : : wt on “a oe
Thorax not so marked .. bid eg av - be ‘i vag LM

. Femora and tibiae with numerous small but conspicuous pale dots ;

abdominal tergites with basal pale bands or spots only quasigelidus, Theo.
Femora and tibiae with the scales mottled, but without conspicuous pale

dots ; abdominal tergites with apical pale bands .. - es ie. - LY)
. Wings with numerous pale scales a ae .. bitaentiorhynchus, Giles.
Wing-scales all dark Mee - ay ais Me .. stmensis, Theo.
. Wings with conspicuous pale markings 7 as - - vias’? JL
Wings unmarked a af ue ae «. IS
. Tip of vein Cu, (lower branch of + ffth) dark- ee 7 .. mimeticus, Noé.
Tip of vein Cu, pale-scaled Fan =f ot a's .. orventalis, sp. Nn.
. Mesonotal scales all dark reddish-brown, except perhaps round the margin ;
middle tibiae without any trace of a pale stripe .. ¢tritaentorhynchus, Giles.
Mesonotal scales mixed light and dark brown ; middle tibiae with a pale
anterior longitudinal stripe more or less indicated = vishnui, Theo.
Femora and tibiae with distinct pale longitudinal stripes anteriorly
(most marked on front and middle legs). . a aia Si eel
Femora and tibia not striped .. af ~ a # 3 gael

Mesonotal scales dark brown, more or less mixed with lighter; pale

abdominal bands generally triangularly produced in the middle
tipuliformis, Theo.

Mesonotal scales reddish-brown ; pale abdominal bands gently rounded
virgatipes, Edw.

Abdominal tergites with complete basal pale bands fe os Sa
Abdominal tergites with basal pale lateral spots only dé os He eA
Abdominal bands white - ag $e ss ivi oe Pen oy)
Abdominal bands pale ochreous oss be oo a aa nh le
Pale abdominal bands very broad; hind tibiae dark except at tip

laticinctus, Edw.
Pale abdominal bands narrow ; hind tibiae with a more or less distinct
pale lateral stripe... ae 2 7 ae .. perexiguus, Theo

330

‘io

20.

10.

11:

12.

13.

14,

15.

NG:

F. W. EDWARDS.

Mesonotal scales ochreous-tinged ie is “a .. _ fatigans, Wied.
Mesonotal scales generally reddish-brown... a0 io .. pipiens, L.
Upper fork-cell in female with a very short stalk; last two joints of
male palpi with a white line beneath ; -. pipiens, Lar
Upper fork-cell in female with a longer stalk : last two joints of male
palpi dark beneath .. a sie an aXe .. laurent, Newst.

Male Hypopygia.

. Side-piece with scales, the lobe scarcely if at all eee the middle,

without flattened plate (Barraudius) 7 vonepee
Side-piece without scales, the lobe well beyond the middle. Be = some

. Clasper long and slender ae ie we o% ne modestus, Fic.
Clasper shorter and stouter.. ; .. pusillus, Macq.

. Clasper with a conspicuous spiny crest ? Ve ees -piece erin a large and con-

spicuous tuft of hairs projecting outwards (Culiciomyia), impudicus, Fic.
Clasper without conspicuous subapical spiny crest ; sea without
conspicuous hair-tuft (Culex)

. Lobe of side-piece without an apically saiated atroned plate ois eo

This plate present, or represented by several. ‘ ot Pepa ys:

. Side-piece with an apical finger-like process ; salapendsees of lobe short

hortensis, Fic.
Side-piece without finger-like process ; appendages of lobe long, apicalis, Adams.

. Lobe of side-piece with several flattened plates a bie hayash, Yam.
Lobe of side-piece with only one flattened plate... a bie Pees 1)

. Plate on lobe of side-piece narrow and pointed as af - si) Ths
The plate broad, rounded, and leaf-like ae a? . a fale

. Tenth sternites without basalarm ... Mt: aguasigelicus Theo.

Tenth sternites with well-developed basal arm

. Mesosome formed of two pairs of upwardly- gece idlae- iaued

structures ; basal arm of tenth sternites short .. bitaeniorhynchus, Giles.
Mesosome formed of one pair of pointed, almost teen structures ; basal

arm of tenth sternites long .. i - .. - stnensts, Theo.
Side-piece with dense hairs round the tip and near the lobe ys A Wl
Tip and region near lobe of side-piece not densely hairy .. ba ot, ee

Clasper greatly widened in the middle, ending in a long, sharp point
orientalis, sp. Ni.

Clasper not much widened in the middle a 4 .. laticinctus, Edw.
Basal arm of tenth sternites well developed .. a kis - Ber x) ee:
Basal arm of tenth sternites very short or absent .. sit Ng Dee te)
Clasper sickle-shaped, gradually tapering to the tip .. Sie Ae ee
Clasper somewhat broadened beyond the middle... se ys oats

Mesosome elaborately divided, the two main divisions each further split ep 15
Mesosome much more simple .. ae Me ors a es see

Lower division of mesosome with only two or three teeth, which are
turned outwards ; .. mimeticus, Noé.

Lower division of mesosome with 4-6 teeth, which are spread out finger-
like or curved tailwards : os e:

Innermost tooth on mesosome considerably longer than the rest
tritaeniorhynchus, Giles.

All the teeth approximately equal in size... oye a vishnut, Theo,

18.

19!

10.

MOSQUITOS OF THE PALAEARCTIC REGION. 331

Mesosome with two divisions, one of which carries two or three short teeth
lipuliformis, Theo.
Mesosome (as seen from above) with three simple divisions .. virgatipes, Edw.
Second division of mesosome simple .. .. perexiguus, Theo.
Second division of mesosome divided into several teeth laurent, Newst.
Second division of mesosome very broad and plate-like .. fatigans, Wied.
Second division of mesosome narrow and hook-like . . ss pipiens, L.

Larvae.

. Siphon rather less than three times as long as broad se ye BZ
Siphon at least four times as long as broad, generally much more eat Weiaes

Siphon pale, all the tufts arranged in a slightly zigzag mid-ventral
line ; tip of antennae black and much narrowed ae pusillus, Mcq.
Siphon blackish ; ventral tufts in three or four pairs, the members of
which are widely separated ; two lateral tufts also present ; antennae

all pale, tip scarcely narrowed es (nebulosus, Theo.|, wmpudicus, Fic.
Comb of eighth segment with 4-8 large sharp teeth - s a: @ nt
Comb of eighth segment with numerous small scales in a triangular patch. . 5
Siphon with a dark ring at one-third of its length; head very dark ;
pecten-teeth 6-9... ays ee ae “ quasigelidus, Theo.
Siphon and head pale ; pecten teeth only 2-3 .. bitaentorhynchus, Giles.
. Siphonal tufts 8-10, in a zigzag ventral row, the first two or three tufts
between the pectens .. .. laticinctus, Edw.
Siphonal tufts more or less paired, none between the. pectens os at AG
Siphon 6-7 times as long as its breadth at the base ; or, if a little shorter
(C. tipuliformis), the pecten teeth have short basal denticles ONY: © aval Abe
Siphon 4—5 times as long as its breadth at the base; pecten teeth with
rather long denticles extending more than half their leneth"—, « Sele

Pecten spines strong, curved, rather wide apart, with small basal denticles

tipuliformis, Theo.
Pecten spines smaller, straight and closer together, generally with more

numerous denticles .. a el ee. os ay ie ee =:
. Siphon distinctly enlarged at the tip, tufts few and small .. apicalis, Adams.
Siphon not enlarged at the tip .. ap at sys Bi . RTs,

. Siphonal tufts rather numerous, some much lenger than the diameter of
the siphon o a <= 0)
Siphonal tufts fewer, none 1e longer than the diameter of the siphon. . ei |

Antennae pale except on the portion beyond the subapical bristles, which
is nearly as long as the part between these bristles and the tuft

mimeticus, Noé.

Antennae ee: or at least more than half, dark, subapical bristles
quite near tip . a5 ws at Ss if as hortensis, Fic.

Antennal tuft at two-thirds; siphonal tufts all subventral (paired)
i tritaentorhynchus, Giles.

Antennal tuft beyond two-thirds; two pairs of siphonal tufts lateral
perexiguus, Theo.
Siphon about 5x 1, pecten teeth averaging 12-15 .. +e pipiens, L.
Siphon scarcely 4x 1, pecten teeth averaging9 .. .. fatigans, Wied.

332 F. W. EDWARDS.

Subgenus Barraudius, nov.

First joint of hind tarsus distinctly shorter than the tibia. No flat scales on top
of head adjoining eyes. Side-pieces of male hypopygium with numerous small
scales on the outer side ; lobe situated scarcely beyond the middle, without flattened
plate, and with only two or three stout spines. Clasper without subapical spiny
crest. Larval mouth-parts normal, not modified for predacity. Siphonal hair
tufts arranged mid-ventrally in a single very slightly zigzag line which runs_ the
whole length. Anal segment short, as in typical Culex. Type species: Culex
pusillus (Macq.), Storey.

Fig. 12. Hypopygia of Palaearctic species of Culex: a, d, f, basal parts, dorsal view; b, ¢, e,
tips of side-pieces, lateral view, ail x 200. a, b, C. modestus, Fic. ; c, d, C. pusillus (Macq.), Storey ;
eaf, C. havashi, Yamada.

The erection of this subgenus is necessary for the reception of two small obscure
species from the eastern Mediterranean region. By several of the characters enumer-
ated above they appear to be more distinct from typical Culex than any other groups
occurring in the Old World.

1. Culex (Barraudius) modestus (T'ic.) (fig. 12, b).

Culex modestus, Ficalbi, Bull. Soc. Ent. Ital. xxi, p. 293 (1890), and xxxi, p. 211
(1899).

Apart from the tarsal character mentioned in the key, the female of this species
is not easy to distinguish from the unbanded variety of C. pipiens. There is, however,
no connection between the two species, which are as widely separated in the structure
of the male hypopygium as any two species of the genus. The average size is smaller

MOSQUITOS OF THE PALAEARCTIC REGION. 333

than the smallest C. pipiens, the integument of the thorax is generally paler and the
scales browner. The long, bare male palpi will at once distinguish that sex from
C. pipiens as well as from all other Palaearctic species except C. pusillus and C. hortensis.
The pale markings of the abdomen have an ochreous tint; the colours of the
tergites are either separated in a straight line, or the pale lateral stripes are slightly
enlarged apically.

Disiribution.—Italy (Ficalbi) ; Hungary (Kertész,; also Neusiedler See, Mik) ;
Macedonia (Waterston) ; Asia Minor (Salyr, Konia and Bashara, Naday); Palestine
(marsh at Tel Abu Zeitun, Austen) ; perhaps the species recorded from Rumania
by Leon as C. fusculus,

WANN

WY
\ WQQn ‘
SS SSS SS y

TERZI

Fig. 13. Culex pusillus (Macq.) Storey, head of larva.

2. Culex (Barraudius) pusillus (Macq.) Storey (figs. 12 c, d, 13, 14).

Culex pusillus, Macquart, Dipt. Exot. Supp. iv, p. 9 (1850).
Culex pusillus, Storey, Bull. Soc. Ent. Egypte 1918, (1919).

I did not at first distinguish this species from C. modestus, and it was recorded
by Barraud under this name. It closely resembles C. modestus, but differs quite
markedly in the male hypopygium. The claspers are shorter and stouter, there
are differences in the lobe of the side-piece, and the anal and genital parts are much
more elongate and rather differently constructed. The pale markings of the abdomen
are pure white, and thus differ in colour as well as in form and position from those
of C. modestus.

I have examined the original specimens of Macquart’s C. pusillus, one of which
is in the Vienna Museum and the rest in the Bigot collection in Mr. Collin’s possession.

(4183) aN

334 F. W. EDWARDS.

All are in such bad condition that they are totally unrecognisable, but from their
size there is no reason to suppose that Storey’s identification is incorrect.

The larva was found by Barraud in small numbers near Basra. The accompanying
figures have been prepared from a comparison of two mounted skins presented by
him to the British Museum. The extremely short siphon, with all the hair-tufts
placed in a slightly zigzag row in the mid-ventral line, is very remarkable, and very
suggestive of the siphon of Luzia, to which genus C. pusillus and C. modestus also

TERZLI ow)

Fig. 14. Culex pusillus (Macq.) Storey, end of abdomen of larva; comb and pecten teeth and
mentum more highly magnified.

show a marked resemblance in the structure of the male hypopygium. There is no.

sign, however, of any modification of the larval mouth-parts for predaceous habits,
and the anal segment is differently shaped.
Distribution.—Egypt (Storey). Mesopotamia (Barraud).

Subgenus Culiciomyia, Theo.

Head in both sexes with a narrow rim of small flat scales along the upper orbital
margin. Male palpi with a row of long scales projecting inwards from the apical

eee ‘ -

MOSQUITOS OF THE PALAEARCTIC REGION. 335

half or more of the long joint ; the scales are of peculiar shape, sharply pointed at
the tip and more or less widened about the middle. Lobe of side-piece of male
hypopygium with two flat plates. Clasper with a conspicuous spiny crest at some
distance before the tip, which is rather suddenly narrowed. Tips of tenth tergites
with the inner spines slender and sharp-pointed, the outer ones much stouter and
blunter. Otherwise as in Culex, s. str.

None of the distinctions mentioned above are fundamental, and it may be doubted
if the subgenus is worth maintaining. The rather well-marked larval characters of
C. nebulosus are not shared by the Oriental members of the subgenus.

One species apparently occurs in the Mediterranean region ; another (C. pallido-
thorax, Theo.) is common in south China, and should be looked for in Japan. There
are other Oriental and Ethiopian species, but none in Australia or America. In the
New World the type is represented by Choeroporpa, Dyar.

3. Culex (Culiciomyia) impudicus, Ficalbi.

Culex impudicus, Ficalbi, Bull. Soc. Ent. Ital. xxii, p. 81 (1890), and xxxi, p. 214
(1899).
(?) Culiciomyia nebulosa (Theobald) Edwards, Bull. Ent. Res. ii, p. 254 (1911).

I do not know Culex impudicus except from Ficalbi’s works, but his figures of
the male hypopygium are so much like the structure to be found in some specimens
of C. nebulosus, Theo., from the Gold Coast* that I feel certain C. impudicus must
be a Culiciomyia closely related to the African species, and it even seems possible
that the apparent differences may be due to inaccuracies in Ficalbi’s figures. This
conclusion is rendered more probable by the fact that I have examined two females
of a Culiciomyia indistinguishable from C. nebulosus in the Paris Museum collection
from Beirut (Dr. Landrieu). The species should therefore be determinable easily
by the characters mentioned under the subgenus. It is an almost uniformly dark
species, with small pale spots at the apical corners of the abdominal tergites.

According to Ficalbi the larvae of C. impudicus were found in water-holes with
much vegetation used for irrigating gardens in Sardinia, and in large marshes in
Sicily. In West Africa C. nebulosus breeds in any small collection of water, especially
about houses ; it is sometimes found in tree-holes and bamboos.

Subgenus Culex, s. str.
(including Neoculex, Dyar).

Head without any small flat scales in the middle in front. Male palpi without a
row of outstanding scales on the long joint. Usually 5-7 pro-epimeral bristles, some
smaller than others. First joint of hind tarsus as long as the tibia, or very slightly
longer or shorter. Side-pieces of male hypopygium without scales, the lobe well
beyond the middle, normally with a flat plate as well as four or five modified bristles.
Clasper without definite spiny crest. Larva with the siphonal tufts more or less
paired, not all in the mid-ventral line, no tufts on basal fourth or more of siphon.

The members of this subgenus exhibit a fair amount of diversity, but it does not
seem to me that any groups are sufficiently circumscribed to be treated as subgenera.
Dyar’s Neoculex (to which belong C. hortensis and C. apicalis) cannot be defined
on any larval character, while even the hypopygial characters on which it is based
are closely approached by C. sinensis, from which species by slight gradations (through
C. quasigelidus and other allied forms) the typical Culex structure is soon reached.

* There are, I find, two definite varieties or species of Culiciomyia in Africa, distinguishable
by hypopygial differences. I have not yet estimated the precise relationship of these two
nor their distribution. If either is synonymous with C. impudicus it will of course have to take
Ficalbi’s name, but it is perhaps more likely that the Mediterranean form is distinct.

(4183) 2a2

336 F. W. EDWARDS.

4. Culex hortensis, Ficalbi.

Culex hortensis, Ficalbi, Bull. Soc. Ent. Ital. xxi, p. 27 (1889), and xxxi, p. 217
(1899).

Maillotia pilifera, Theobald, Mon. Cul. iv, p. 274 (1907).

Culex geniculatus, Theobald (nec Olivier), Mon. Cul. iii, p. 216 (1903).

Easily distinguished from the other Palaearctic species with dark tarsi by the
apically situated bands on the abdominal tergites and from its ally C. apicalis by
the white spot at the tip of the hind tibia, and the bare male palpi. The abdominal
bands are variable in width, being reduced occasionally to lateral spots only.

The larvae are said to prefer weedy ponds, particularly those covered with duck-
weed.

Distribution.—Throughout the Mediterranean region and central Europe, extending
as far north as Paris and Berlin. Some new records are: Corsica (Mann); Asia
Minor (Ereckli, Sabanja, v. Bodemeyer); Germany (Berlin, Schildhorn, Oldenberg,
19); Transcaspia (Firudza, C. Ahnger); Syria (Beirut, Landrieu).

5. Culex apicalis, Adams.

Culex apicalis, Adams, Kansas Univ. Sci. Bull. ii, p. 26 ( ? June 1903).

Culex sergenti, Theobald, Mon. Cul. iii, p. 218 (July 1903).

Culex pyrenaicus, Brolemann, Ann. Soc. Ent. France, Ixxxvii, p. 427 (1919).
Culex territans, Howard, Dyar & Knab, Monogr. iv, p. 293 (1912) (nec Walker).

In spite of the great differences in the male hypopygium, there can be no doubt
that this is closely related to C. hortensis. The most obvious distinctions of
C. apicalis are the hairy terminal joints of the male palpi and the dark tip of the hind
tibia. The wing-scales seem to be a little narrower, and the pale abdominal bands
are also perhaps on the average narrower. It may not always be possible to dis- -
tinguish the females with certainty ; Eckstein states that they differ from those of

C. hortensis in having the bases of the abdominal sternites dark-scaled, but I cannot
_ confirm this.

The larva differs from that of C. hortensis in the bicoloured antennae, in the
shape of the siphon, and in the smaller and less numerous siphonal tufts. The two
species are said by Séguy to breed under similar conditions, though according to
Eckstein C. apicalis is found in clear water.

Distribution.—Occurs over a wide area in Europe and North America, apparently
also in North Africa, though I have seen only females from there (including Theobald’s
type of C. sergenti) and am not absolutely certain of their identity. Some new records
are: Tunis (Tamerza, Langeron, 9); Italy (Gorizia, Mik, 3 2) ; Carniola (Wippach,
Handlirsch) ; Transcaspia (Amudaria, C. Ahnger).

6. Culex hayashi, Yamada (fig. 12 e, f).
Culex hayashi, Yamada, Dobuts. Z. Tokio, xxix, pp. 61-72 (1917).

This differs from all other species known from the Palaearctic region in having
the male palpi straight and considerably shorter than the proboscis (about three-
quarters as long), but there are several other Oriental species with which it might
be confused, such as C. brevipalpis (Giles) and C. jenseni (Meij.). The male
hypopygium is also very distinct, on account of the structure of the mesosome and
the numerous plates on the lobe of the side-piece ; in the former point C. hayashi
much resembles the subgenus Lophoceratomyia, but it does not show any modification
of the male antennae, nor any flat scales on the top of the head ; it should perhaps
be placed in Dyar’s subgenus Neoculex,if that is adopted. The species is unicolorous
brown, only the lower side of the abdomen somewhat lighter. The scaling is that
of a normal Culex, but there seem to be some flat scales on the prothoracic lobes.

“sy

MOSQUITOS OF THE PALAEARCTIC REGION. 3o7

Dr. Lamborn found the larvae in muddy pools in company with those of
Anopheles punctibasis.

Distribution.—Japan (Tokio, Yamada, a series presented by the collector to the
British Museum in 1915; Nagasaki, Lamborn).

7. Culex quasigelidus, Theobald.

Culex quasigelidus, Theobald, Mon. Cul. iii, p. 181 (1903) ; Edwards, Bull. Ent.
Res) it pyeos Work)

This is one of the most distinct members of a rather large group of tropical species,
which Theobald included in his genus Leucomyia. The leg markings are distinctive,
but are not at all unlike those of the Old World species of Lutzia, especially L. tigripes,
of which C. quasigelidus has been taken to be a variety, though in reality it is very
different. The larvae, like many others with long siphons, live in weedy pools ;
they are very similar in structure to those of the other members of this group.

Distribution.—Widely spread in the Ethiopian region, occurring in Madagascar,
and spreading northward by the Nile valley as far as Alexandria.

8. Culex bitaeniorhynchus, Giles.

Culex bitaeniorhynchus, Giles, J. Bombay Nat. Hist. Soc. xiii, p. 607 (1901) ;
Edwards, Bull. Ent. Res. iv, p. 231 (1913).

The wing-scales of this species are unusually broad for a Culex, and on this
account Theobald placed it in the genus Taentorhynchus, with which it has really
no connection. Usually the pale scales on the wings are almost as numerous as the
dark ones, at least in the female, but a variety occurs in which they are comparatively
few and scattered. The femora and tibiae are also very much mottled. The pale
bands of the abdomen are very variable in width ; they may be very narrow, or the
abdomen may be almost all pale. The species is semi-domestic, the larva often living
in polluted water.

Distribution.—Throughout the Oriental region ; occurring also in Japan and North
Australia. A variety, differing slightly in the male hypopygium, is widely spread
in Africa.

9. Culex sinensis, Theobald.

Culex gelidus var. sinensis, Theobald, Mon. Cul. iii, p. 180 (1903).
Leucomyia sinensis, Theobald, Mon. Cul. v, p. 313 (1910).
Culex sinensis, Edwards, Bull. Ent. Res. iv, p. 231 (1913).

This is at first sight very much like C. bitaeniorhynchus, but differs in the much
narrower and entirely dark wing-scales, and very considerably in the male hypopygium.
Apart from this, the femora and tibiae are less mottled, the pale scales which are
present tending to be aggregated into small dots, though these are not nearly so
conspicuous as in C. bitaeniorhynchus.

Distribution —Widely spread in the Oriental region, and, like the last species,
occurs also in Japan (Tokio, Yamada), but is not known from Australia or Africa,
where it seems to be represented by allied but distinct species.

10. Culex mimeticus, Noé.
Culex mimeticus, Noé, Bull. Soc. Ent. Ital. xxxi, p. 240 (1899).

A very interesting species on account of the spotted wings, the markings com-
prising three pale ochreous areas on the costa, which extend on to the first vein,

338 F. W. EDWARDS.

also other pale areas, the most noticeable of which are in the middle of the third vein
and towards the base of the sixth. The fifth vein is entirely dark, except for a part
of its upper branch. The wing-scales are narrow but rather short. The proboscis
has a well-defined pale ring about the middle in both sexes. The male palpi have
pale rings at the bases of the last two joints and a very narrow one at the tip of the
last joint. The side-pieces of the hypopygium are only moderately hairy ; the lobe
with the usual five modified bristles (the apical one unusually flattened and outwardly
directed) and leaf-like plate; clasper sickle-shaped, gently tapering, with well-
marked terminal claw ; tenth sternites with the basal arm quite long, though shorter
than the sternites ; second division of mesosome split into two or three teeth.

The larva has been partly described and figured by Martini, but he has omitted
to notice an important point, the position of the pair of subapical antennal bristles
only a little more than mid-way between the tuft and the apex of the shaft. It is
also noteworthy that the pale colour of the antenna extends some way beyond the
tuft, almost to the subapical bristles. The 4 or 5 pairs of.larger siphonal tufts are
at least twice as long as the diameter of the tube. The larva is remarkably like that
of C. hortensis, differing in antennal characters and in the rather greater number
of siphonal tufts; the latter point is probably not of much importance, as the
number and also the position of the tufts is certainly variable in many species of the
genus.

The most interesting fact about this species is its occurrence in association with
Anopheles superpictus, to which it bears a considerable resemblance in wing-markings.
Whether we have a genuine case of mimicry, and if so what advantage the species
could gain by it, I will not attempt to judge, but it is perhaps an even more remarkable
fact that the allied C. mimulus, which differs in having a dark third vein, occurs with
Anopheles culicifacies or A. minimus, which differ in the same way from A. superpictus.

Distribution.—Mountainous regions in the eastern Mediterranean region. Italy
(Noé); Macedonia (Martini, Waterston); Palestine (Cropper); Cyprus (Miss
Bate).

The species was till recently supposed to have a much more extended distribution,
but I have recently shown that the form inhabiting Ceylon and Malaya differs slightly
both in wing markings and hypopygial details, and have therefore treated it as
a distinct species, C. mimulus. There are good larval differences between C. mime-
ticus and C. mimulus, the latter having few and short siphonal tufts, and the subapical
antennal bristles close to the tip. A second form, which is probably equally distinct,
is found in Hong Kong, Formosa, and South India (Ootacamund, recorded by me
recently as C. mimeticus). This differs from the true mimeticus in the much broader
pale tip to the longer male palpi, the absence (apparently not quite constantly) of
the basal arm of the tenth sternites, and perhaps in other details. I have seen only
females from North India, and cannot say whether they belong to this second Oriental
form or to the true mimeticus. The Japanese form must obviously be treated as
another quite distinct species.

11. Culex orientalis, sp. n.

Differs from C. mimeticus as follows :—Wing-scales somewhat broader and
distinctly ionger, the wings therefore appearing more densely scaled. Cu, (lower
branch of fifth vein) with a pale area at its tip, most noticeable in the female ;
another pale area (more or less developed) before the fork. Male proboscis with
numerous pale scales on the apical portion beyond the ring, sometimes the whole
apical portion is pale. Hypopygium: side-pieces large and stout, densely hairy,
especially round the somewhat produced tip and near the lobe; lobe with eight
somewhat flattened appendages, all much alike, with rounded, not hooked tips,
and placed almost in a continuous row ; besides these there are numerous accompany-
ing long hairs ; leaf-like plate and its accompanying bristle present as usual. Clasper

MOSQUITOS OF THE PALAEARCTIC REGION. 339

very large, flat, very much broadened a little beyond the middle, ending in a rather
long sharp point ; terminal claw very minute ; subapical spiny crest slightly indicated.
Tenth sternites with moderate basal arm. Second division of mesosome with three
rather large teeth and about five small ones.

The hypopygium is more like that of C. laticinctus than that of C. mimeticus.
It would be of interest to know whether there is any similar resemblance in the larvae.

Distribution.—Japan (Tokio, Yamada). <A series presented by the collector
to the British Museum in 1915 was determined by me then as C. mimeticus, but a
closer study reveals the striking differences enumerated above. Also Yokohama and
Kobe (Lamborn) ; the larvae in rice-fields in company with Anopheles hyrcanus.

12. Culex tritaeniorhynchus, Giles.

Culex tritaeniorhynchus, Giles, J. Bombay Nat. Hist. Soc. xiii, p. 606 (1901) ;
Edwards, Bull. Ent. Res. iv, p. 233 (1913), and vii, p. 224 (1917).

This species is sufficiently distinguished by the characters mentioned in the
key, but I have given a number of others in the papers quoted above. The average
size is very small (3 mm.), but in this respect the species varies a good deal in different
parts of its range, Japanese examples being much larger than those from Palestine.

Larvae have been received from Capt. Barraud from Mesopotamia; they are
remarkably similar to those of C. perexiguus, described below ; I can discover very
few differences beyond those mentioned in the key, which seem most likely to be
constant. Other larvae from Ceylon differ slightly from these, but not to such
an extent that they need be separated specifically. They are found usually in
salt marshes, often in company with other small species with a banded proboscis
(C. vishnut or C. sitiens).

Distribution.—Palestine and Mesopotamia (Barraud) ; Palestine (IKhirbet Hardrah,
Austen ; Jerusalem, Goldberg); Japan (Tokio, Yamada ; Chuzenji, Gallots ;
Nagasaki, Lamborn) ; China (Shanghai, Lamborn, etc.). Also throughout the Oriental
region and on both the east and west coasts of Africa.

13. Culex vishnui, Theobald.

Culex vishnut, Theobald, Mon. Cul. i, p. 355 (1901) ; Edwards, Bull. Ent. Res. iv,
p. 233 (1913), and vii, p. 225 (1917).

In spite of the very slight difference in the hypopygia (there are perhaps some other
slight distinctions besides the one mentioned in the key) I feel sure this species is
distinct from C. tritaeniorhynchus, its closest ally and frequent associate. C. vishnue
breeds in rice-fields, salt-marshes, and elsewhere. No isolated larvae have been
received at the British Museum, nor has.a description of the early stages been
published. The hypopygium is very similar to that of C. mimulus.

Distribution.—Mesopotamia (Barraud) ; Japan (Osaka, Theobald). Also through-
out the Oriental region, but as yet unknown from Africa.

14. Culex tipuliformis, Theo. (figs. 15, 16).

Culex tipuliformis, Theobald, Mon. Cul. ii, p. 325 (1901) ; Edwards, Bull. Ent.
Res. ii, p. 262 (1911), and iti, p. 31 (1912).

Culex creticus, Theobald, Mon. Cul. iii, p. 189 (1903).

Apart from the striped femora and tibiae, and the more or less produced abdominal

bands, this might easily be mistaken for C. pipiens, especially in rubbed specimens.
It is, however, generally darker in colour, the upper fork-cell is not so long, and the

340 F. W. EDWARDS.

cross-veins, though variable in position, tend to be more approximated than in
C. pipiens, being occasionally almost in one line. As in C. pipiens, the last two joints
of the male palpi have whitish markings beneath, but in this species the pale scales
tend to be arranged more in patches, one of which is at the tip of the last joint.

The larva has been described by Bedford from the Transvaal (U.S. Afr. Dept.
Agr., 5th & 6th Repts. Director Vet. Res., 1919, p. 741), his description and
figure agreeing in the main with specimens I have examined, though he shows
shorter and more numerous tufts on the siphon. The accompanying figures are
based on Capt. Barraud’s material. The siphon is distinctly longer than that of
C. pipiens, index about 5-5-6.* The pecten teeth are 6-9 in number, but rather
widely spaced, and reaching beyond a third of the length of the siphon. The
first few teeth are quite small, but the last four or five are long, curved, and almost
simple, only one or two small basal denticles being present. The antennae are
dark at the tip and at the extreme base, pale in the middle, the tuft being placed at
about three-fifths. The head is more or less extensively dark basally. The siphon

Fig. 15. Culex tipuliformis, Theo., head of larva.

is generally all pale, but among the specimens sent by Capt. Barraud from Mesopotamia
there are several which have the basal half of the siphon dark or even black ; these
specimens also appear to have the siphon a little shorter than usual, but the adults
issuing from them do not differ appreciably from normal C. tipuliformis.

Distribucion.—Atlantic islands; Mediterranean region generally; extending
through East Africa to the Cape and by way of Persia into north India and Assam ;

* By the siphonal index I mean the ratio of the diameter of the base of the siphon to the
length, the valves not being reckoned into the length. Séguy apparently takes the ratio of
the average width to the length.

MOSQUITOS OF THE PALAEARCTIC REGION. 341

a rather remarkable distribution, which is exactly parallelled by that of Theobaldia
longiareolata. I believe the following are new records :—Asia Minor (Konia and
Bashara, Naday) ; Persia (Enzeli, Buxton) ; West Caspian (Lenkoran, Karsch) ;
Libyan Desert (Bulag, W. J. H. King).

Fig. 16. Culex tipuliformis, Theo., end of abdomen of larva; comb and pecten teeth and
mentum more highly magnified.
15. Culex virgatipes, Edw.
Culex virgatipes, Edwards, Bull. Ent. Res. v, p. 126 (1914).

Differs from C. tipuliformis in the male hypopygium, which is extremely similar
to that of the African C. trifilatus, Edw., in the uniformly reddish-brown mesonotal
scales and in the abdominal markings. The cross-veins are on the average more

342 F. W. EDWARDS.

widely separated than in C. tipuliformis, and the femoral and tibial stripes, though
quite distinct, are rather narrower in this species. The resemblance to C. pipiens |
is much more close than in the case of C. tpuliformis, the leg-markings affording
the main external distinction. The larva is undescribed; it was found by Dr.
Lamborn in company with that of C. fatigans.

Distribution.—So far as our present knowledge goes, this species is confined to
the eastern part of the Palaearctic and Oriental regions, into which C. tipuliformis
apparently does not extend. Hong Kong (Macfarlane) ; Sikkim (Wyville- Thompson) ;

Vladivostok and River Amur (Wuorentaus :, Helsingfors Museum); Shanghai
Lamborn).

16. Culex laticinctus, Edw.
Culex laticinctus, Edwards, J. Proc. Asiatic Soc. Bengal, ix, p. 49 (1913).

This species differs markedly from C. pipiens, the one to which it approaches
most nearly in size and appearance, in the pure white abdominal bands, which are
as broad as or broader than the dark bands which alternate with them. The male
palpi are no longer than the proboscis, and are less hairy than those of C. pipiens.
The prothoracic lobes often show a number of flat scales on the lower part. The
hypopygium is not unlike that of C. orientalis, especially in the form and hairiness
of the side-piece, but the appendages of the lobe and the structure of the mesosome
are different.

The rather remarkable larva has been described and figured by Storey as “ Culex
sp. no. 2258.”’

Distribution.—Throughout the Mediterranean region ; Canary Islands (Orotava,
Graham-Smith) ; southern Spain (Aguilas, G. Boag) ; southern France, and as far
north as Paris (Ségwy) ; Tunis (Tamerza, Langeron; Djerba, in coll. Bezzt) ; Ana-
tolia (Budrum, Mus. Civ. Genova); Cilicia, Syria and Palestine (Barraud; Mt.
Carmel, Austen ; Jerusalem, Goldberg) ; Egypt (Storey) ; Arabia (Muscat, Gill).

17. Gulex perexiguus, Theobald (figs. 17, 18).
Culex perexiguus, Theobald, Mon. Cul. i, p. 199 (1903).

The very small size of this species, together with the narrow white (not ochreous-
white) abdominal bands, and the pale stripe on the outer side of the hind tibia (not
always very clearly marked, and in the male sometimes indistinguishable), will
serve to separate it from other members of the group with dark tarsi and basally
banded abdominal tergites. Apart from this, and the aedoeagal structure, C. perext-
guus may be known by the colour of the mesonotal scales, dark brown mixed with
brassy ochreous.

I cannot detect any difference whatever between the adults of C. perexiguus
and the West African form of C. wnivittatus, either in external characters or male
hypopygial structure. The larvae, however, seem to be utterly different. Some
confusion has existed regarding the larvae of C. univittatus, but Dr. Ingram
assures me that the larva described and figured by him and Dr. Macfie (Bull. Ent.
Res. x, p. 68) was identified by the isolation method, and he is sure that no error
occurred. The figure indicates a larva similar in many respects to that of C. quast-
gelidus, but with several remarkable features, such as the possession of only a single
pair of minute siphonal tufts and an incomplete ring on the anal segment.

Larvae of C. perexiguus sent from Palestine by Capt. Barraud differ in practically
every detail from the larva described by Ingram and Macfie, almost the only point

MOSQUITOS OF THE PALAEARCTIC REGION. 343

of resemblance being in the length of the siphon. The accompanying figures have
been prepared from isolated skins sent by Capt. Barraud, the adults issuing from
which I have examined. The following is Capt. Barraud’s description :—

“Antenna light in colour except towards base and tip. Shaft clothed with
spicules. Antennal tuft of about 24 subplumose hairs arising at about three-quarters
from the base. Mid frontal hair tufts of subplumose hairs; ante-antennal tuft
of 8 hairs; outer median tuft of 2, inner median of 3. Small lateral tuft above eye
of about 4 very small hairs. Mental plate with 7 teeth on either side of the central
one, the outermost tooth some distance below the others.

‘‘ Siphon about seven times as long as the width at base. Pecten of from 11-14 teeth;
teeth slightly curved, with three secondary spines on one side ; last few teeth slightly

Fig. 17. Culex pevexiguus, Theo., head of larva.

more detached than the remainder. Hair tufts on siphon represented by about four
pairs of very short and fine hairs (towards tip usually single). Tufts on eighth segment
of from 4 to 6 subplumose hairs. Comb of small teeth in triangular patch. Anal
gills about the length of the anal segment, the dorsal pair rather longer than the
ventral. Two or three hairs in the tuft on the dorsal edge of the analsegment. Brush
well developed, about 12 tufts each with about 6 hairs.”

Since it is impossible to consider two such different larvae as belonging to the
same species, there is no alternative but to revive Theobald’s name ferexiguus for
the Mediterranean form.

Distribution.—Palestine (Cropper, Barraud). Since it is impossible to separate
the adults, the further distribution of C. pevexiguus as distinguished from C.
univittatus cannot be given, but it seems reasonable to assume that the form is the
same throughout the Mediterranean region. From an exarhination of adults and
from Storey’s remarks on the larva it is obvious that this is the species he has recorded

344 F. W. EDWARDS.

from Egypt as C. decens ; the larva of the African C. decens is indeed very similar,
though the hypopygium differs. It is perhaps also the species recorded from
Algeria by the Sergents as C. fatigans, since they state that the siphon is longer than
that of C. pipiens. Adults, probably of C. perexiguus, have been received from
southern Spain (Fowler), Muscat (Gill) and Amritsar, Punjab (Barraud).

Fig. 18. Culex perexiguus, Theo., end of abdomen of larva ; comb and pecten teeth and mentum
more highly magnified.

18. Culex laurenti, Newst.

Culex laurentt, Newstead, Ann. Trop. Med. i, p. 24 (1907); Edwards, Bull.
Ent. Res. v, p. 70 (1914).

The female of this species is almost impossible to distinguish from the unbanded
variety of C. pipiens.* The male differs from both C. pipiens and C. perexiguus
in having no pale line beneath the last two joints of the palpi. I have figured the

MOSQUITOS OF THE PALAEARCTIC REGION. 345

hypopygium in the paper quoted. According to Storey the larvae are indistinguish-
able from those of C. perexiguus.

Distribution An Ethiopian species, known from Madagascar, Zanzibar, and
the Congo, which has also been recorded from Egypt by Storey as C. tmvidiosus.
Also found in Egypt by Austen (Kantara, Suez Canal). Perhaps occurs in Palestine
(Acre and Jerisheh, Awusten ; females only).

19. Culex fatigans, Wied.

Culex fatigans, Wiedemann, Aussereurop. zweifl. Ins. p. 10 (1828).
Culex quinquefasciatus (? Say), Howard, Dyar and Knab, Monogr. iii, p. 345
(1915).

This species may be distinguished from C. pipiens by the combination of some
or all of the following characters :—Mesonotal scales somewhat coarser, with a
dull brownish-ochreous instead of a dark brown or reddish-brown tinge (but
Japanese and American specimens are often as red-tinged as C. pipiens). Upper
fork-cell shorter in both sexes, that of the female being less instead of more than
three times as long as its stem. Male palpi somewhat shorter and less hairy. Pale
abdominal bands of the female rather more rounded. The only absolutely reliable
distinction between the adults is, however, in the structure of the aedoeagus (see
figures in Bull. Ent. Res. iv, pp. 54, 55). The larvae are rather more easily separated
than the adults by the characters mentioned in the key.

Distribution.—This species, almost universal in the tropics, has a very limited
area of occurrence within the Palaearctic region. I have only seen it from Lower
Mesopotamia (Barraud), Seistan, eastern Persia (Annandale), and Japan (Kobe and
Nagasaki, Lamborn). It has been recorded by various observers from southern
Europe and North Africa, but I consider it highly probable that all such records
refer to other species.

20. Gulex pipiens, L.

Culex pipiens, Linnaeus, Syst. Nat. Ed. x, p. 602 (1758).

(2) Culex fasciatus, Miiller, Fauna Insectorum Fridrichsdalina, p. 87 (1764).

(?) Culex molestus, Forskal, Descriptiones Animalium, p. 85 (1775).

(?) Culex luteus, Meigen, Klass. i, p. 6 (1804).

(?) Culex domesticus, Germar; Reise nach Dalmatien, p. 290 (1817).

Culex rufus, Meigen, Syst. Beschr. i, p. 7 (1818).

(?) Culex bicolor, Meigen, Syst. Beschr. i, p. 9 (1818).

(?) Culex pallipes, Waltl, Reise Tyrol etc. ii, p. 110 (1835).

Culex pallipes, Macquart, Dipt. Exot. i, i, p. 33 (1838).

(?) Culex pallipes, Meigen, Syst. Beschr. vii, p. 1 (1838).

(?) Culex meridionalis, Leach, Zool. Journ. ii, p. 292 (1825).

Culex marginalis, Stephens, Zool. Journ. i, p. 455 (1825).

(?) Culex thoracicus, Robineau-Desvoidy, Mém. Soc. d’Hist. Nat. Paris, ili, p. 409
(1827).

(?) Culex calcitrans, Robineau-Desvoidy, loc. cit.

(2) Culex rufinus, Bigot, Expl. Scient. Tunisie Dipt. p. 7 (1888).

Culex agilis, Bigot, Ann. Soc. Ent. France (6) ix, Bull. cxii (1889).

Culex phytophagus, Ficalbi, Bull. Soc. Ent. Ital. xxi, p. 126 (1890), and XXVIi1,
p. 286 (1896).

Culex haematophagus, Ficalbi, Bull. Soc. Ent. Ital. xxv, p. 143 (1893).

Culex pallens, Coquillett, Proc. U.S. Nat. Mus. xxi, p. 303 (1898).

Culex melanorhinus, Giles, Gnats, p. 342 (1900).

Culex longifurcatus, Becker, Mitt. Zool. Mus. Berlin, ii, p. 68 (1904).

346 F. W. EDWARDS.

Culex nigritulus, Theobald, Mon. Cul. i, p. 140 (1901) (nec Zetterstedt).

Culex varioannulatus, Theobald, Mon. Cul. iii, p. 198 (1903).

Culex azoriensis, Theobald, Mon. Cul. iii, p. 210 (1903).

Culex quasimodestus, Theobald, Ann. Mus. Nat. Hung. iii, p. 88 (1905).

Culex osakensis, Theobald, Mon. Cul. iv, p. 439 (1907).

Culex pipiens var. doliorum, Edwards, Entom. xlv, p. 263 (1912).

(2?) Culex nigritulus, Wesenberg-Lund, Danske Vid. Selsk. Skr. Nat. Math. Afd.

(8) vii, p. 131 (1921).

The distinctions in external characters between the adults of C. pipiens and
C. fatigans have been enumerated above. I have found them reliable in sorting out
Mesopotamian specimens, where the two species occur together and C. pipiens
is fairly constant ; but in the southern and eastern Mediterranean region C. pipiens
is subject to so much variation that the hypopygial differences would have to be
relied on; the structure of this organ in C. pipiens is fairly constant, the slight
variation that does occur showing usually little or no approach to C. fatigans.

The examination of a number of mounts of hypopygia of Japanese specimens,
however, seems to show that the Japanese race of C. pipiens differs constantly from
the European, having the second division of the mesosome much broader than usual
and the third division not quite so stout, thus being to some extent intermediate
between C. pipiens and C. fatigans. Theobald’s type male of Culex osakensis, which
in 1912 I took to be C. fatigans, belongs to this form, but Coquillet’s name pallens
is no doubt also applicable, and should be used to designate the variety. It would
seem from their figures that Dyar and Knab’s Culex comitatus, described from
California, belongs to this var. pallens rather than to typical C. pipiens, andit is quite
likely to have been introduced into California from Japan. In both C. pipiens and
C. fatigans a minute basal arm to the tenth sternites may be present or absent.

A variety of frequent occurrence in the Mediterranean region has the pale bands
of the abdomen reduced to lateral spots, either in the female only, or in both sexes.
There are also two other varieties worthy of special mention. In North Africa
many specimens occur with the mesonotal scales more or less ochreous, and in some
the dark parts of the abdomen also tend to this colour, so that it is tempting to
assume that we have here an incipient modification in colour to suit desert conditions.
I have examined the hypopygium of one specimen so coloured, and have no doubt
as to its identity. Theobald has described this variety as C. quasimodesta, but it
is doubtful if it is at present more than a sporadic variation. The second variation
is in the length of the upper fork-cell of the female. Over the greater part of the
range of the species this is fairly constant, but in the Levant and Asia Minor many
specimens are found in which the cell is shorter than usual, and little, if any, longer
than that of C. fatigans. It seems rather significant that this very region is on the
borders of the range of C. fatigans ; the possibility of interbreeding may be indicated,
but, against this, it should be noted that such intermediate specimens have not been
found in Mesopotamia, where the two species are known to occur together.

I am indebted to Capt. Barraud for calling my attention to the existence of
what seems to be a definite larval variety of C. pipiens in Palestine and Syria. In
this form the average number of pecten-teeth is 12, and there seems to be little varia-
tion from this; of 39 specimens critically examined by Capt. Barraud, no fewer
than 24 had either 12 or 13 pecten teeth, the number in the remaining specimens
varying from 9 to 17. Further distinctions of this Levantine race are the smaller
average size, the lighter-coloured antennae, and the shorter average length of the
siphon (index about 4-5).

Capt. Barraud found that in Mesopotamian specimens the average number of
pecten teeth was greater and the range of variation more. Out of 27 specimens
examined the average number of teeth was 15°7; only 10 specimens had either 15

MOSQUITOS OF THE PALAEARCTIC REGION. 347

or 16 teeth, the number in the others ranging from 12 to 20. These specimens also
had dark antennae, and the average size was larger and the average length of the
siphon rather greater (index about 5). I find that specimens from Britain and
Macedonia, though rather variable, agree in the main with this Mesopotamian type,
which may therefore be taken as the common [uropean form.

I have not been able to detect any constant difference between Palestine adults
and those of other countries. Both the banded and unbanded forms occur there,
and, as mentioned above, some (but by no means all) of the females have the upper
fork-cell shorter than usual. It may be noted that in its several peculiarities the
Palestine larva of C. pipiens approximates to that of C. fatigans.

It is possible, as long ago suggested by Ficalbi, that there are two races of this
species, differing little, if at all, externally, but one being more addicted to sucking
human blood than the other. In England C. pipiens will certainly attack man at
times, but can seldom be regarded as troublesome ; I have never myself experienced
its bite, nor found a blood-gorged female in a bedroom. In south Europe, however,
the reports of various observers lead one to suppose that it is more regularly addicted
to feeding on human blood. Further experience may possibly show that the
Palestinian type of larva described above is widely distributed in the Mediterranean
region and represents the more troublesome race. If this should be proved to be
the case the varietal name molestus, Forskal, might be applied to this form.

Wesenberg-Lund describes as C. migritulus, Theo., a Culex larva which seems
to differ in many respects from C. pipiens: e.g., in the shape of the mentum and
of the pecten-teeth and comb-scales and in the longer siphon. As I have not seen
the adults reared from these larvae, I will only remark that the larvae of my C. pipiens
var. doliorum (which I considered identical with Theobald’s C. migritulus, and which
I do not now consider even varietally distinct from C. pipiens) conform fairly well
to Wesenberg-Lund’s description of C. pipiens.

Synonymy.—It is impossible to say what species were actually intended by
most of the old descriptions, but I think it probable that the names C. bicolor, Mg.,
C. pallipes, Mg., C. thoracicus, R.-D., C. calcitrans, R.-D., and perhaps also C. luteus,
Mg., were based on more or less rubbed specimens of this species. From the habits
indicated by Forskal and Germar for C. molestus and C. domesticus it seems probable
that this species was intended, C. fatigans being excluded owing to its now apparently
established absence from Europe and Egypt. The description of C. pallipes, Waltl,
was evidently supplied by Meigen, and amplified by him in 1838. The British
Museum possesses a copy of Meigen’s Abbildung eur. zweifl. Ins., hand-coloured by the
author, in which the figure of C. rufus evidently represents C. pipiens, though the
venation is shown in a conventional manner.* In his diagnosis of C. meridionalls,
Leach says ‘“‘ abdomine segmentis omnibus postice griseo marginatis,’’ but as he makes
a similar statement regarding his C. micaensis and C. musicus, it seems probable
that by “ postice’’ he meant “ basally.’”’ Ficalbi’s description of C. phylophagus,
especially as regards the male palpi and abdominal bands, shows that he had
C. pipiens, not C. laticinctus or C. univittatus, before him. I have examined the
types of C. marginalis, C. agilis, C. varioannulatus, and C. azoriensis, and find
them to be C. pipiens. Dr. Dyar informs me that he has examined Coquillett’s
tvpe of C. pallens, and that it is C. piprens. The species has frequently been
referred to as C. ciliaris, L., but I think probably incorrectly.

Distribution —Throughout the Palaearctic region ; also in parts of North and
South America, East and South Africa, and Madagascar; no doubt spread by
commerce.

* It may also be remarked here that the figures in this work of C. vevans and C. annulipes
agree with the interpretation of these names adopted in this paper. Some of the other figures
are less decisive.

348 F. W. EDWARDS.

Bibliography.

The following list includes only the more important of the works dealing with
Palaearctic CULICIDAE from the entomological point of view; purely economic
works, as well as obsolete systematic papers, are not included. A fairly full biblio-
graphy, up to the end of 1919, is given by Martini in “ Ueber Stechmiicken.”’

Barraup, P. J. Notes on some Culicidae collected in Lower Mesopotamia.—Bull.
Ent. Res. x, 1920, pp. 323-325.

——. Mosquitos collected in Palestine and adjacent territories.—Bull. Ent. Res.
xi, 1921, pp. 387-395.

Biackiock, B. and Carter, H. F. Observations on Anopheles (Coelodiazesis)
plumbeus, Stephens, with special reference to its breeding-places, occurrence in
the Liverpool district, and possible connection with the spread of malaria.—
Ann. Trop. Med. xii, 1920, pp. 413-446, pls. x—xil.

BROLEMANN, H. W. Sur quelques Culex de Pyrénées et description d’une éspéce
nouvelle.—Ann. Soc. Ent. France, Ixxxvii, 1918, pp. 425-440; Ixxxvii, 1919,
pp. 65-103; Ixxxix, 1920, pp. 51-73. (Includes good figures of the male
hypopygia of most of the species found in France.)

Carter, H. F. Descriptions of the male genital armatures of the British Anopheline
mosquitoes.—Ann. Trop. Med. xiii, 1920, pp. 453-457 figs.

CuristopuErs, S. R. The male genitalia of Anopheles.—Ind. J. Med. Res. iu,
1915, pp. 371-394, pls. xx—xxv.

—_—. Notes on some Anophelines from Arabia and Mesopotamia.—Ind. J. Med.
Res. iii, 1915, pp. 180-200, pl. xvi.

———. Recent observations on the Anopheles of the Middle East.—Ind. J. Med.
Res. vii, 1920, pp. 710-716, 3 maps.

and Suortt, H. E. Malaria in Mesopotamia.—Ind. J. Med. Res. viii, 1921,
pp. 508-552.

Eckstein, F. Zur Systematik der einheimischen Stechmticken—Centralbl. f. Bakt.,
Parasit. u. Inf.-Krankh., 1 Abt., Ixxxii, 1918, pp. 57-68, figs. (females) ; Ixxxin,
1919, pp. 281-294, figs. (larvae) ; Ixxxiv, 1920, pp. 223-240, figs. (males).

_._ Die einheimischen Stechmiicken. Eine Schilderung ihrer Lebensweise und
anleitung zu ihrer Bestimmung, Mit 17 Abbild.—Wiss. Mitglied des
Forschungsinst. f, angew. Zool. Berlin, 1920, 58 pp. (Largely a reprint of above
series of papers, with some biological and other details added.)

FicaLsBi E. Revisione delle specie europee della fam. delle Zanzare.—Bull. Soc.
Ent. Ital. xxiv, 1892, pp. 257-284; xxv, 1893, pp. 48-61, 136-144; xxvi,
1894, pp. 66-75, 315-320 ; xxvii, 1895, pp. 29-38; xxviii, 1896, pp. 108-313,
pls. icv. (The earlier sections include quotations of the original descriptions
of all European species then known.)

_ Venti specie di Zanzare (Culicidae) Italiane.—Bull. Soc. Ent. Ital. xxxi, 1899,
pp. 46-234, figs.

Forry, H. Etude morphologique de Pyretophorus chaudoyer Théob., aux differents
stades de son evolution—Campagne antipaludique de 1911, Alger, 1912, pp.
49-50, pls. ili-v.

GArtLL-VALERIO, B. Sechzehn Jahre Untersuchungen iiber Kuliziden und Malaria.
—Arch. Schiffs. Hyg. xxii, 1918, pp. 154-158. (Briefly summarises the author’s
work, with bibliography of his 45 short papers.)

GOETGHEBUER, M. Culicides et Corethrides de Belgique-——Ann. Soc. Ent. Belg.
1910, pp. 81-87, 410-412.

MOSQUITOS OF THE PALAEARCTIC REGION. 349

Joyeux, C. Note sur les Culicides de Macédoine.—Bull. Soce bath i xot..ih 191s:
pp. 530-547.

Lance, W. D. Handbook of British Mosquitoes.—London, British Museum (Natural
History), 1920.

LANGERON, M. Remarques sur les larves du Culex geniculatus et sur les larves
de Culicinés pourvues d’un long siphon.—Bull. Soc. Path. Exot. ix, 1916,
pp. 438-442, figs.

——. Remarques sur l’évolution larvaire de Theobaldia annulata (Schrank, 1776).—
Bulls Socw Path: Exot. ix; 1916; pp. 703-708, figs.

—. Morphologie et biologie de la larve de Theobaldia spathipalpis, Rondani,
1872.—Bull. Soc. Path. Exot. xi, 1918, pp. 98-103, figs.

——. La larve d’ Anopheles chaudoyet (Theobald, 1903).—Bull. Soc. Path. Exot.
xi, 1918, pp. 291-297, figs.

Leon, N. Studii asupra Culicidelor din Romania.—Bucuresti, 1910, Directiunea
Generale a serviciului sanitar. Pp. 274, pls. xv, figs. 111.

——. Contributions 4 l'étude des Culicides de Roumanie.—Centralbl. f. Bakt.
liii, 1910, pp. 499-505.

Martini, E. Uber Stechmiicken, besonders deren europaische Arten und ihre
Bekampfung.—Archiv f. Schiffs- und Tropenhyg. Band 24, Beiheft 1, 1920,
pp. 1-267, pls. i-v, 117 text figs.

—. Die biologische Malariabekampfung in Mazedonien.—Zeitschr. f. Angew. Ent.
vii, 1921, pp. 225-286.

DE Me1jeErE, J. C. H. Zur Kenntnis niederlandischer Culiciden—Tijd. v. Ent.
liv, 1911, pp. 137-157, pls. vii-x.

Nutratt, G. F. H. and SuipLey, A. E. The structure and biology of Anopheles.—
Journ. Hyg. I, 1901, pp. 45-77, 269-276, 451-482, pls. i, ii, viii-x; IL, 1902,
pp. 58-84, 166-215, pls. vi-viii.

SCHNEIDER, P. Beitrag zur Kenntnis der Culiciden in.der Umgebung von Bonn.—
Verh. Nat. Ver. Preuss. Rhein]. u. Westf. Ixx, 1914, pp. 1-54, 2 pls.

Skécuy, E. Les moustiques de France.—Bull. Mus. d’Hist. Nat. Paris, 1920,
pp. 51-58, 141-149, 223-230, 322-329, 407-414, 512-519.

——. Remarques sur quelques larves de moustiques—Bull. Soc. Ent. Fr. 1920,
pp. 309-311.

——. Note sur l’armure génitale du Culex jugorum Villen.—Bull. Soc. Ent. Fr.
1921, pp. 39-40. .

SERGENT, Ed. et Et. Observations sur les moustiques des environs d’Alger.—Ann.
Inst. Pasteur, xvii, 1903, p. 60.

——. Observations sur les Anopheles d’Algerie.—Ann. Inst. Pasteur, xix, 1905,
p. 144, and xx, 1906, p. 393.

Storey, G. Keys for the determination of Egyptian Mosquitoes and their larvae.—
Bull. Soc. Ent. Egypte, Cairo, 1918 (1919), pp. 84-106, 2 pls.

VASSILIEV, J.B. Cellia pulcherrima, Theo., und einige neue Angaben tiber Erscheinen
und Uberwintern von Anopheles bifurcatus, L., in Turkestan. (See Rev. App.
Bate, wp. 193.) if

WatTERSTON, J. On the mosquitos of Macedonia.—Bull. Ent. Res. ix, 1918, pp. 1-12.

WESENBERG-LuND, C. Anatomical description of the larva of Mansonia richard
(Ficalbi) found in Danish freshwaters.—Vidensk. Medd. Dansk. naturhist. Forn.
Ixix, 1918, pp. 277-328, figs.

—. Contributions to the biology of the Danish Culicidae——D. Kgl. Danske
Vidensk. Selsk. Skrifter, Nat. Math. Afd., 8 Raekke, vii, 1921, pp. 1-210,
pls. i-xxi.

(4183) 2B

350

F. W. EDWARDS.

Index of Specific and Varietal Names applied to Palaearctic Mosquitos.

(Synonyms and unrecognisable names in italics.)

Page
abfitchit, Felt .. 25/305
aegypti, L. ae 73 O20
affinis, Steph. .. .. 288
agilis, Big. : Wo O40
alaskaensis, Ludl. -. 288
albescens, sp. n. .. 308
albionensis, Macegr. go oael0)

albolstevalis, Theo. Ao acoults}
albopalposus, Becker .. 325
albopictus, Skuse setae

albopunctatus, Rond. .. 319
algeriensis, Theo. 2. 20
alineata, Schn. .. OS
alpinus, L. .. 309
angustealatus, Becker a2 O20
annulatus, de Fourc. .. 288
annulatus, Be 3.8 Asts)
annulatus, Schrank. .. 288
annulipes, Mg. .. yo mills)
antennatus, Becker So SHA
apicalis, Adams .. 336
avabica, Giles .. ve 299
avabicus, Becker sig PASS)

avabiensis, Patton (A) .. 277
avabiensis, Patton (C) .. 322

ayvcanus, Blanch. be BOUL
arctica, Edw. .. ye vaste
articulatus, Rond. ee:
azoriensis, Theo. .. 346
bambusa, Yam. 56 4syil
bergrothi, Edw. .. Aid Aoi
bicolor, Mg. a .. 345
biturcatusy ls. Se SOFA
bimaculata, Leic. eeeeo
bipunctatus, R.-D. 3 O07
bitaeniorhynchus, Giles 337
broquettii, Theo. .. 300
calcitvans, R.-D. OLD
calopus, Mg. .. .. 325
cantans, Mg. Ai .. 304
cavdamitisi, Newst. dio) PAIRS
casplus, Pallas .. 50 Zag)
cataphylla, Dyar 2 OLO
chaudoyei, Theo. 7 200
christophi, Port. .. 285
ciliaris, L. sis Bh ere:
cinereus, Mg. .. -« 323
claviger, F. a6 Se ae
claviger, Mg. see DATE
cleopatrae, Willcocks 3 280)
communis, Deg. .. 34
CONCLINNUS, Steph. a Oe
costalis, Theo. .. Bi ATI
creticus, Theo. .. Sip) cores)
cretinus, sp. n. .. 56. OPA
Currver, \COGan he). .. 300
cyprius, Ludl. .. ap OOM
detritus) Elals) =: ms xe co)

diantaeus, H.D.K. ace ey It

diplolineata, Schn.
diversus, Theo. ..
doliovum, Edw.
domesticus, Germar
dorsalis, Mg. ..
dorsovittatus, Villen.
a@thali, Patton ..

eatoni, Edw.
echinus, Edw.
elegans, Fic.
elutiisSpemiae ae
excrucians, Walk.

fatigans, Wied. ..
fasciatus, F.
fasciatus, Mg. ..
fasciatus, Miller
ficalbii, Noé
flavescens, F. ..
flavescens, Miller
jlavescens, Theo.
flavirosiris, Mg.
flavovirens, R.-D.
flavus, Motch.
flerowii, Portch.
fletcheri, Coq.
freyi, sp. n.
fumipennis, Steph.
fusculus, Zett.

gallii, Mart.
geniculatus, Ol...

glaphyropterus, Schin. ,

gvahami, Ludl. ..
gvisescens, Steph.
guttatus, Curt.

haematophagus, Fic.
haplolineata, Schn.
hargreavesi, Edw.
hayashi, Yam. .
hispaniola, Theo.
hortensis, Fic.
hyrcanus, Pall. ..

impiger, Walk. ..
impudicus, Fic.
impunctus, Don.
innuttus, D. & K.
intrudens, Dyar

japonicus, Theo.
jugorum, Villen.

konoupi, Brullé
koreicus, Edw. .

Page

. 315
.. 308
.. 346

. 345
.. 300

~ 3h2

5 arks}

1 OO
.. 320
2. O20
ot ge afle)

. 305

-. 345
DO OAS)
. 314

». 320
. 318

lateralis, Mg. .. ollie

laticinctus, Edw. « 342
laurenti, Newst. .. 344
lepidonotus, Edw. .. 308%)
lesnet, Séguy .. 312
leucacanthus, Lw. Us
leucogrammus, Lw. no ey)
leucomelas, Mg. .. .. 314
leucopygus, Eysell. . 323
lewist, Ludl. : . 272)
lindesayi, Gilesier 7 203
longiareoiatus, Macq. .. 287
longifurcatus, Beck. .. 345
longisquamosa, Theo. .. 299
lukisi, Chr. 5 47K)
/uteovittatus, Theo. 55 wohlls)
lutescens, F. 2. 306
luteus, Mg. . 345
macedoniensis, C. & H... 278
maculatus, Mg. . .. 304
maculicosta, Beck. ote, eat
maculipennis, Mg. Baie
maculiventris, Mcq. . 300
malariae, Grassi + 322
marginalis, Stph. se (O40)
MATIAe, SLSeen wae 3 O08
maroccanus, d’Anfr. .. 287
mauritianus, Grp. -. 279
meigenanus, Dyar .. 313
melanorhinus, Giles oo 345
mervidionalis, Leach .. 345
mesopotamiae, Chr. . 274
metalepticus, Dyar . 316
mimeticus, Noé.. BO ore
modestus, Fic. .. Au prep
molestus, Forsk. y 340
morsitans, Theo. .. 289
multicolor, Camb. So Aske)
musicus, Leach . 308
myzomyfacies, Theo. .. 279
nearcticus, Dyar . 309
nebulosus, Theo. He carey)
nemorosus, Mg... 3813, 314
nicaensis, Leach. 50 PAsts
nigrvina, Eckst. .. a OZ
nigvipes, Staeg. 50 Pail
nigvipes, Zett. . 309
nigrvitulus, Zett. a. OZ
nigritulus, W.-L. .. 347
niveus, Eichw. ae OLE
niveus, Ludl. sor coulis}
nursel, Theo. SS
obscurus, Mg. Te Oke
obturbans, Walk . 326
occidentalis, D: & K. 2. 272
onondagensis, Coq. . 300
orientalis, sp. n. .. 338
ornatus, Mg. .. a5 OL
osakensis, Theo. .. 346

Page
319

pallens, Coq. .
palestinensis, Theo.
pallipes, Mcq.
_pallipes,Mg. ..
pallipes, Waltl. ..
palmeni, Edw. ..
paludis, Theo.
parvulus, sp. n.
parvus, Mcq. :
penetrans, R.-D.
penicillaris, Rond.
perexiguus, Theo.
persicus, var. n.
pharoensis, Theo.
phytophagus, Vic.
pictus, Lw. s
pilifera, Theo.
pipiens, L. ee
plumbeus, Steph.
prodotes, Dyar .
pseudonivea, Theo.
pseudopictus, Grassi
pulcherrimus, Theo.
pulchripalpis, Rond.
pulchritarsis, Rond.
pullatus, Coq.
punctatus, Mg. ..
punctibasis, sp. n.
punctor, Kirby ..
pungens, R.-D. ..
pusillus, Macq. ..
pyrenaicus, Brolem.

quadratimaculatus, Macq. 308

quartus, Mart.
quasigelidus, Theo.

MOSQUITOS OF THE PALAEARCTIC REGION.

Page

.. 346
6 PAINS)
.. 345
. 345
.. 345
5 old
son ao
.. 314
sep o22
Se ashe)
5 oath
. 342
. 280
ee PALE
.. 345
.. 274
.. 336
. 345

quasimodestus, Theo.

Page

. 346

quinquefasciatus,H.D.& 1.345

veptans, Mg. ate
rhodesiensis, Theo.
richiardii, Fic.
vostochiensis, Mart.
rufinus, Big.
vufus, Mg.

vufus, Gimm.
rusticus, Rossi ..

sacharovii, A.
salinellus, Edw.
salinus, Fic.
scutellaris, Theo.
selengensis, Ludl.
semicantans, Mart.
sergenti, Theo. (4)
sergenti, Theo. (C)
servvatipes, Beck.
sevus, Mart.
siberiensis, Ludl.
siculus, R.-D.
sinensis, Theo. ..
sinensis, Wied. ..
spathipalpis, Rond.
stephensi, Liston
sticticus, Mg. ..
subalbatus, Coq.
subochrea, Edw.
subtilis, Serg.
sugens, Theo.
superpictus, Grassi
surcouft, Theo. ..

sylvae, Theo.
sylvaticus, Mg.

terviet, Theo.
tervitans, H. D. K.
theobaldi, Meij. ..
thoracicus, R.-D.
tipuliformis, Theo.
titillans, Leon
togoi, Theo.
towadensis, Mats.
trifurcatus, F. .
tritaeniorhynchus,

unguiculata, Edw.
univittatus, Theo.

310, 311

351

Page

sole
. 315

336

-« 290
. 345
. 339

pee AS

-. 318
. 285

271

Giles 339

.- 283
. 342

variegatus, Schrank. 288, 396

varioannulatus, Theo. ..
= 267;
ae O22
7 Qa
se Oa
Asi
nine Ketohe
P7325
eye yAeL
2 267

vassilievi, Portch.
vexans, Mg.
villosus, R.-D.
virgatipes, Edw.
viridis, R.-D.
vishnui, Theo.
vittatus, Big.
vorax, sp. n.
vulgaris, L.

waterhouset, Theo.
willcocksi, Theo.

zammtitii, Theo.

346

. 304
AS)

. 303

353

AGRILUS FOVEICOLLIS, MARS., AS A CAUSE OF THE DECAY
OF THES CULTURE ,OH “ROSES IN BULGARIA.

By Pror. S. A. MoKRZECKI.
Experimental Agricultural Institute, Sofia.

During the months of May and June 1921 I investigated the cause of the serious
losses that had occurred among the rose-trees (especially Rosa damascena, Mill.)
which are extensively grown in Bulgaria for making attar of roses. The conclusion
arrived at is that the cause of the general decline in rose culture is inadequate

Galls on stems of Rosa damascena, caused by the larvae of Agyvilus foveicollis,
Mars.—1, One-year-old shoot ; the bark is cut off and the burrows of the
three to four days’ old larva may be seen; no swelling is formed yet.
2, Two-year-old shoot, with a swelling near its base; the top of the shoot
was beginning to dry up. 3, Two-year-old shoot, with the swelling cut
open; several annular passages are clearly visible. 4, Three-year-old
shoot, on which the swelling is very distinct ; by this time the majority
of the small branches had already faded. All photographs taken from
nature and slightly enlarged.

354 PROF. S. A. MOKRZECKI.

nourishment of the plants, owing to an insufficient amount of humus and nitrogen
in the soil, aggravated by the entire neglect of manuring by the growers. But
the immediate cause of the death of thousands of rose-bushes has proved to be
certain galls that are to be found on the stems, reaching 2-3 cm. in length and
sometimes twice as thick as the normal stem.

When the gall is cut open, dark-coloured burrows may be seen circling the stem
just under the bark, the number varying from three to fourteen. Up till now,
however, no insects have ever been found in these galls, and previous investigators
attributed them to Agvilus viridis, L.* Injuries of this kind have been also recorded
on roses in Italyt and France,t and Houard§ attributes them to the activity
of larvae of some Microlepidopteron.

My investigations have led me to the conclusion that the galls are caused by
the larvae of Agrilus foveicollis, Mars.,|| a species described from Siberia and never
recorded previously from any locality in Europe.

The life-cycle of this new and very destructive pest of roses is not quite clear,
but the following points have been actually observed by myself. The beetles appear
in the middle of May and live on the leaves of roses, nibbling their margins. The
female, after copulation, lays her eggs, up to 30 in number, each separately, under
the bark of one-year-old shoots. Each egg is laid in a scarcely perceptible oblong
hole, made by the ovipositor ; it is about 0-3-0-4 mm. long, white, with the surface
covered with a network of fine furrows. The eggs hatch after 5-7 days, and even
during the first 2-3 days of its life the larva may make as many as three annular

* burrows around the shoot, under the bark, filled with black excrement. No swelling,
however, is formed during the first summer (fig. 1). In the second year the swelling
is noticeable and increases gradually (figs. 2 and 3), while the infested shoot begins
to dry up. In the third year the whole stem gradually dies (fig. 4).

The life of the larva lasts apparently about one year, and that is why larvae
are not to be found in the galls which are already well formed. There may be several
galls on the same stem.

* Dr. Nikoloff, M. Stefanoff and N. Pouchkareff : ‘‘ The Culture of Roses in Bulgaria,’’ Revue
d’Inst. des Recherches agronomiques en Bulgarie, i, nos. 5 & 6, 1921, pp. 11-14 (in Bulgarian).
There is a figure of A. viridis, L., and its larva and cocoon, with descriptions taken from
Richter von Binnenthal’s book: ‘‘ Die Rosenschaddlinge aus dem Tierreiche,”’ Stuttgart (1903),
as well as original figures of the burrows as observed by the authors in Bulgaria.

+ Del Guercio, ‘‘ Intorno ad una deformazione del fusto della Rosa in Italia.”,—Nuove Relaz.
Staz. Entom. Agr. Firenze, pp. 143-146, pl. ix, figs. 1-2.

t J. Beauderie, ‘‘ Les Croussins du Rosier.”—Hort. Nouv. Lyon, 1911.

§ ‘‘ Les Zoocecidies des plantes d’Europe,”’ i, p. 542.

|| Coleopt. Hefte, v, 1869. I am much indebted for the identification of my specimens to
the Imperial Bureau of Entomology and to its Director, Dr. G. A. K. Marshall.

355

NOTES ON A COLOUR TROPISM OF ASTEROCHITON (A LEURODES)
VAPORARORIUM, WESTWOOD.

By Ln. Lroyp, \se-"(Leeds).

During 1919-20 an investigation into the habits of Asterochiton ( Aleurodes)
vaporariorum, Westw., was being carried out at the Lea Valley Experimental Station,
Cheshunt, with a view to controlling its attacks on tomatoes under glass. The
station exists for the study of scientific problems connected with the glasshouse
industry, and as the staff is small, the work is necessarily confined to strictly economic
lines. Points that appear in the course of the investigations cannot be followed to
their ultimate conclusion in these earlier years of the station’s existence if they seem
to be of theoretical rather than of practical importance. The following notes

relate to such a problem, which it has been decided to record, though in a very
unfinished state.

A habit of this whitefly of settling upon clothing led to a short study of its colour
reactions. All the experiments were carried out in a heated greenhouse in December
and January. A wooden cage was used, 18in. by 18in. by 24 in. high, with the
bottom six inches forming a wooden well, and above this four sides and a roof of
muslin. An exactly similar wooden framework without the muslin was also
employed, its sides and roof being open to the greenhouse.

Experiments with Transmitted Light.

Various weak solutions in water of substances conveniently to hand were made
up. Clear white glass corked tubes one inch in diameter and three in length were
filled with the solutions to a depth of one and a half inches, and the outsides of the
tubes to the same depth were thinly smeared with a clear adhesive composed of resin
and castor oil. The tubes were then hung by strings around the four sides of the
frame so that they were equidistant from each other and about fifteen inches from
the centre of the well. Cut infested foliage was placed in the floor of the well. During
the tests the positions of the tubes were interchanged, the least attractive to the
position of the most attractive, and so on, or the whole frame was revolved through’

90°. The insects were removed by means of a needle and counted at intervals of
from one hour to two days.

Experiment 5.

December 3rd; bright sun; exposure, seven hours of daylight. Twelve weak
solutions were made up at random to give a wide range of colours ; 452 flies were
trapped, and their distribution is shown in diagram 1 by the continuous line. The
following solutions were attractive in order of merit: light yellowish-green (picric
acid+-jodgriin), fluorescent (weak eosin), greenish-yellow (picric acid), orange
(orange G.). The following gave negative results : water, bright red (fuchsin), blood

red (Congo red), blue-green (methyl green), pale blue (methylene blue), indigo
(nachtblau) and violet (gentian).

Experiment 6.

December Sth-6th; dull; exposure 16 hours’ daylight. Solutions as above,
with three others added ; 266 flies were trapped, and their distribution is shown
in diagram 1 by the broken line. The same solutions as in the last experiment,
together with purple (potassium permanganate), proved negative. The eosin solution
was only moderately attractive, anda strong solution of potassium bichromate rather

356 Tt, LLOYD:

less so, but a weak solution of the same substance was the most attractive in the
test, trapping nearly a quarter of the total flies caught.

It was evident from these and from other crude tests that the attractive solutions
all had a predominating yellow colour, or were fluorescent. The next experiments
were carried out in an endeavour to find an optimum attraction.

Experiment 9.

December 9th—15th ; little weak sunlight ; exposure 56 hours’ daylight. Solutions
of potassium bichromate (yellow fluorescent) of percentage strengths 5, 1, 4 7g
at zh two tubes of each ; 1,771 flies were trapped, and their distribution: is shown
in diagram 2. The optimum strength for this solution was clearly from 4 to 3's per
cent., from the nature of the curve between these two; +4 per cent. potassium
bichromate was therefore used as a standard in the next experiment.

Percentages of total 4/es
°

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alle Cae i ages) Oh Ls eae 88

vd > v9 c 9 gi he ox Pa VS % Ga ve

3 GN) Uo ueee e kee oe Ee ES oS Syncs
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Diagram 1. (See Experiments 5_and 6.)

Experiment 10.

December 16th-20th; little weak sunlight ; exposure 40 hours of daylight.
Four solutions of orange G. of strengths }, 35, 74s, and 54, per cent., and four of eosin
of strengths 1, 4, 4, and z4, per cent., one tube of each, were tested against four
control tubes of + per cent. potassium bichromate ; 983 flies were trapped, and had
the distribution shown in diagram 3. The eosin solutions were relatively unattractive,
but the nature of the curve shows that there was some attraction, the optimum
strength being about ;4, per cent. It may be pointed out that the occasion when
the attraction of this substance was powerful was when tested in bright sunlight,
its other tests being all in dull weather with intermittent weak sun. The solutions
of orange G. were less attractive than the control except the z+, per cent. solution,
which drew 60 per cent. more flies. In the course of the experiment three counts
were made on the twelve tubes, and this tube trapped most flies in each case, except
that it was bettered once by a control, 53 as against 42. An attempt was made to
analyse the maximum point on the orange G. curve in the next experiment.

NOTES ON A COLOUR TROPISM OF ASTEROCHITON VAPORARIORUM, WESTW. 3097

Experiment 12.

December 20th—23rd ; little weak sunlight ; 32 hours’ daylight. Three strengths of
orange G., ;, 7, zis per cent., four tubes of each ; 849 flies were trapped as follows:
iz per cent., 257 flies (30 per cent.) ; zs percent., 274 flies (32 per cent.) ; ote per
cent. 318 flies (37 percent.). This is not a decisive result, although the figures appear
to be graded, the differences being within the range of experimental error and rather
erratic in the separate counts.

The solution which drew forth the greatest response was thus one of orange G.,
of a strength approximating to 7}, per cent.

175

150

se 125

2 100

Percentages of tote/ Fi1es
Percentages of Flies th relation 0 contso/

TAS)
15 50
10 25
Ss

“, Ys Y2e Ysi2

Orange & feonrtinuous line) \ Percentage
Ya Ye Yea Yese SEPEAg Cue

| zl | i !
a a a a Pe iy /256 Losin (broken sine)
ercentage strength of solution 1°, ; .
¥ Pot 2 ; Yah Potassium bichromate
° otassiulm brchromate contro/ [ea*ted sine)
Diagram 2. (See! Experiment-9;) Diagram 3. (See Experiment 10.)

Experiments with Reflected Light.
Experiment 3.

December 2nd ; a strip of white card, seven inches long and one and a half inches
broad, was divided by transverse lines into seven equal sections, and these were
painted with water-colour washes in the following order: red, orange, yellow, green,
blue, indigo, violet. When dry this was lightly smeared with a transparent adhesive
and placed horizontally in the cage one foot above the bottom of the well, which
contained infested foliage. The flies left the foliage and flew to the top of the cage,
and were then in a position to see the trap. The card was exposed for two hours,
the sun being bright. At the end of the first hour the card was turned, to reverse
the position of the colours. In all, 397 flies were trapped, and their distribution is
shown in diagram 4. Orange, green and yellow proved attractive, the last-named
trapping 51 per cent. of the total. The four remaining colours held only about
10 per cent. of the total catch.

Experiment 11.

December 19th—20th ; a small tomato leaf one and a half inches long was taken,
and six pieces of paper of the same size and shape were washed with water-colours
as follows: red, orange, yellow, yellowish-green, blue and white. Each of these

(4183) 2c

358 LL. LLOYD.

was then fastened in sandwich manner between a pair of white glass microscope
slides and the upper one was lightly smeared with adhesive. These were placed in
the cage as described above and their positions were interchanged twice, counts.
being made each time. The total exposure was four hours, with the sun shining
brightly ; 1,041 flies were caught, and their distribution is shown in diagram 5.
Yellow trapped 60 per cent. of the total; the yellowish-green was attractive, and
the tomato leaf less so. Red, blue, white, and in this case orange also, gave negative
results.

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Diagram 4. (See Experiment 3.) Diagram 5. (See Experiment 11.)

The large numbers trapped in these short exposures by the opaque yellow, com-
pared with the numbers trapped by the yellow solutions, suggests that the former
1s more attractive, and the following experiments seem to confirm this.

Experiment 13.

_ December 22nd—29th ; five glass tubes twelve inches long and half an inch in
diameter were taken. Four were filled with a solution of ;{; per cent. orange G., which
in a column of this diameter appeared to have the same tint as the zs per cent. solution
ina one inch column. The fifth tube was lined with yellow paper. All were smeared
with adhesive and suspended around an infested tomato plant ten inches high on
staging in the greenhouse. The clear tubes trapped 178 flies, an average of 44 each,
and the opaque one 91. At the end of the experiment there remained on the plant
17 flies only.

The relative attractions of tomato foliage and a yellow surface are shown in the
following experiments. ; ;

Experiment 14.

December 23rd-31st; a sheet of yellow paper, 7 in. by 8 in., was sandwiched
between two sheets of glass of the same size and the trap was coated with adhesive on
both sides. This was then suspended vertically facing north and south, and a lightlv
infested tomato plant about one foot in height was placed on each side of it at a
distance of a foot. A third similarly infested tomato plant was placed at a distance
of six feet. All the plants were recently infested, and no flies emerged on them during
the course of the experiment. There were, however, a number of infested plants
in the greenhouse on which flies were emerging, and these had access to the three
plants and the trap. Eight days after the experiment had been set up the plant
to the north held 33 flies and the north side of the trap 602, the plant to the south

NOTES ON A COLOUR TROPISM OF ASTEROCHITON VAPORARIORUM, WESTW. 359

held 21 and the south side of the trap 470. The distant plant held 122 flies. The
trap was now removed, and ten days later the two plants which were near it and were
kept in the same position held 80 and 133 flies respectively, no emergence having
occurred upon them and the source of invading flies remaining approximately the
same as when the trap was exposed.

Experiment 16.

January 8th ; the main source of flies was a nettle plant growing in a pot and
massively infested. A small uninfested tomato plant one foot in height was placed
at a distance of two feet and the tube lined with yellow paper, as described in
experiment 13 above, was stuck into the soil of the pot holding the tomato. After
four-and-a-half hours of moderate sunlight a count of the flies was made. The
tomato plant with a total foliage area (both surfaces of leaves) of 120 sq. in. held
106 flies, about one to the square inch, and the trap with an area of 14 sq. in. held
160 flies, or about 11 to the square inch.

Conclusions.

Both sexes of the insect were drawn to the yellow traps, so the habit is probably
not associated with mating. The bodies of the mature insects are yellow and the
scale has a large internal organ of a yellow colour, the mycetoma, which is con-
spicuous in the younger scales. The insect is distinctly gregarious, quite apart from
the fact that it seeks younger foliage, and sometimes the majority of the flies on a
plant are found gathered on one or two leaves. It appeared possible that this gre-
garious habit was at the root of the attraction, and a number of adults were enclosed
in a glass tube. This and an empty tube were smeared with tanglefoot and placed
in the cage. The tube of fly caught 28 and the empty tube 25 in eight hours’
exposure. There was clearly no definite attraction here, and it is probable that the
yellow body of the insect is not sufficiently conspicuous to be perceived.

Several manifestations of the phenomenon were observed apart from the experi-
ments recounted. The attraction to clothing, and especially to the khaki garments
which were commonly worn by the tomato hands at this time, has been referred to.
The flies may often be seen on yellow flowers. It was noticed particularly that
the yellow flowers of narcissus were more favoured than the foliage of the plant,
and many eggs were laid on them, though these inevitably perished. In the autumn,
when heavily infested tomato houses were being cleared out in the neighbourhood,
and the air in places was full of the disturbed insects, great numbers wereseen hovering
over yellow privet, but the insect does not care for leathery foliage and has not been
known to breed on this shrub.

The most favoured foods of the insect out of doors in England are the following
plants :—hollyhock, calceolaria, tobacco, sunflower, French and runner beans, and
vegetable marrow. All these have a distinctly vellowish-green foliage. The colour
attraction is believed to be associated with choice of food and to draw the migrating
insects to light-foliaged plants as opposed to those which have dark leaves; but for
the reason given, the elaborate experiments which would be necessary to prove
this could not be undertaken. It is possible also that a solution of the problem could
only be found by studying the insect in its native haunt, which is said to be Brazil.

It might be possible to devise some method of partial check te the pest by means
of these traps. Literally enormous numbers can be caught on a yellow screen by
shaking infested plants and holding the screen near them. The flies fly up when
disturbed in this way and pass to the screen as though drawn by a magnetic force.
Any such method of check would be clumsy and is not worth elaboration, since
perfect control of the pest can be obtained in the greenhouses by fumigations with
hydrocyanic acid, and out of doors, in England at any rate, the insect is not a pest
of major importance, and can only rarely be considered a pest at all.

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:

COLLECTIONS RECEIVED.

The following collections were received by the Imperial Bureau of Entomology
between Ist July and 30th September 1921, and the thanks of the Managing
Committee are tendered to the contributors for their kind assistance :—

Dr. W. M. Apers, Government Economic Biologist :—75 Culicidae, 53 Hippo-
boscidae, 4 other Diptera, and 6 Coleoptera ; from Zanzibar.

Mr. E. BALLARD, Government Entomologist :—7 Ants and 157 Rhynchota ;
from South India.

Mr. C. F. C. BEEson, Forest Zoologist :—28 Braconidae and 47 Curculionidae
from India.

Mr. G. E. Bopktn, Government Economic Biologist :—77 Culicidae, 1 Tabanus,
4 Hymenoptera, and 1 Beetle; from British Guiana.

Prof. C. K. BRAIN :—4 Culicidae, 3 Haematopota, 25 other Diptera, 105 Coleoptera,
41 Rhynchota, and 101 Orthoptera ; from Stellenbosch, South Africa,

Mr. P. A. Buxton :——58 Diptera, 14 Hymenoptera, 50 Coleoptera, 6 Rhynchota
and 81 Orthoptera ; from Palestine.

Prof. T. D. A. CocCKERELL :—28 species of Coccidae ; from Colorado.

Mr. J. B. Corporaat, Entomologist, Algemeen Proefstation, Medan :—8
Rhynchota ; from Sumatra.

Mr. R. Corram :—12 Cockroaches; from the Anglo-Egyptian Sudan.

Department of Agriculture, Bangalore :—G64 Coleoptera ; from India.

Mr. T. BAINBRIGGE FLETCHER, Imperial Entomologist :—232 Coleoptera, and

1,505 Orthoptera ; from India.

Mr. E. CRESSWELL-GEORGE :—4 Coleoptera and 4 Orthoptera ; from Nyasaland.

Mr. C. C. GowpveEy, Government Entomologist :—8 Lvyperosia, 1 Stomoxys, 6
Hymenoptera, 16 Coleoptera, 42 Lepidoptera, 5 species of Aphididae, 12 species
of Coccidae, 25 other Rhynchota, and 1 Dragonfly ; from Jamaica.

Mr. C. B. HARDENBERG, Chief Entomologist, Department of Agriculture :
1,268 Coleoptera ; from Portuguese East Africa.

Mr. H. HarGreaAves, Government Entomologist :—2 Hippoboscidae, 53 other
Diptera, 37 Siphonaptera, 411 Hymenoptera, 118 Coleoptera, 600 Thysanoptera,
15 Lepidoptera, 24 species of Coccidae, 76 other Rhynchota, 1 Acridiid, 70 Anoplura,
14 Mites, 64 Ticks, 3 Spiders, and 2 Pseudoscorpions ; from Uganda.

Hawaiian Sugar Planters’ Association :—36 Coleoptera ; from Honolulu: and
66 Curculionidae and 4 microscope preparations ; from Australia.

Mr. G. F. Hirt, Entomologist, Australian Institute of Tropical Medicine :—
64 Culicidae and 4 Tabanidae; from Australia.

Dr. A. INGRAM :—21 Tubes containing 700 minute Diptera ; from the Gold Coast.

Dr. W. B. JoHNSON :—5 Culicidae, 1 Chrysops, 3 Hippocentrum, 71 Haematopota,
11 Tabanus, 25 Glossina, 3 pupa cases, and a species of Mite attacking Glossina ;
from Northern Province, Nigeria.

Mr. H. H. Kino, Government Entomologist :—13 Gryllid nymphs associated
with Ants, 3,513 Ants and a number of early stages; from the Anglo-Egyptian
Sudan.

(4183) 2D

362 COLLECTIONS RECEIVED.

Dr. W. A. LAMBoRN :—980 Culicidae and 30 other Diptera ; from China’ and ‘
Japan.

Rev. T. P. Levetr :—2 Chalcididae ; from Richmond, Yorkshire.

Dr. J. W. Scorr MAcrig :—282 Culicidae, 19 Psychodidae, 1 Tabanus, 1 Glossina,
1 Asilidand prey, 700 other Diptera, 10 Ants, 12 other Hymenoptera, 18 Coleoptera,
34 Lepidoptera, 2 species of Coccidae, 101 other Rhynchota, 8 Cockroaches and 1
nymph, 16 Planipennia, 1 Dragonfly, 6 Anoplura, 2 Spiders, a number of Mites,
2 Scorpions and young, and 2 small Snakes ; from Accra, Gold Coast.

Mr. C. W. MALLy, Senior Entomologist, Department of Agriculture, Cape Town :—
5 Curculionidae ; from South Africa.

Mr. G. E. Mertor.:—2 Wasps; from the Anglo-Egyptian Sudan. |

Ministry of Agriculture, Egypt :—41 Hymenoptera, 98 Coleoptera, 1 Moth, 1
Reduviid bug, and 3 Orthoptera; from Egypt.

Prof. S. A. Moxrzeck! :—8 Agrilidae ; from Bulgaria.

Mr. J. C. Moutton :—49 Coleoptera, 1 Pentatomid bug, and 17 Orthoptera ;
from Singapore.

Dr. L. PERINGUEY, Director, South African Museum :—94 Coleoptera; from
South Africa.

Senhor A. F. DE SEABRA:—I18 Coleoptera, 9 Lepidoptera, and 1 Ascalaphid ;
from San Thomé.

Mr. H. W. Simmonps:—1 Tabanus, 25 other Diptera, 21 Hymenoptera, 8
Coleoptera, 10 Lepidoptera, 11 Lepidopterous pupa cases, | Caddisfly, 2 species
of Coccidae, 2 other Rhynchota, 1 Cricket, 1 Haemerobiid, 2 Spiders, 5 Mites, and
1 Micro-slide of Nematode eggs; from Fipi.

Dr. R. J. Trrryarp, Chief, Biological Department, Cawthron Institute of
Scientific Research :—3 species of Aphididae ; from New Zealand.

Mr. F. W. Urtcu, Government Entomologist :—10 Coleoptera, 12 Pentatomidae,
and 150 Acarina; from Trinidad.

Mr. Ropert VEITCH :—13 Diptera, 61 Hymenoptera, 222 Coleoptera, 47 Lepidop-
tera, 85 Rhynchota, 12 Orthoptera, and 7 Odonata ; from Fiji.

Mr. H. S. WALLACE :—Examples of corks destroyed by Lepidopterous larvae ;
from Newcastle-on-Tyne.

Mr. O. H. Watters :—125 Coleoptera; from the Simla Hills, India.

Prof. H. H. WuetzeL, Department of Agriculture, Bermuda :—5 Coleoptera ;
from Bermuda.

He ma AN

xl. Part 4p. 363 84. Res.
| FEBRUARY, 1922.

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363

ONT SOME AUSFRALIAN TERMITES OF* THE’ GENERA
DREPANOTERMES, HAMITERMES AND LEUCOTERMES.

By GERALD FP. Hin, FES:
(Plates IX-—XII.)

This paper is intended as the first of a series in which it is proposed to record
the results of an examination of a large collection of termites from various parts of
the Australian continent. The greater part of this collection has been gathered
in the northern districts of the Northern Territory, and in North Queensland, but
during the past year it has been considerably increased by the addition of numerous
small collections received from correspondents in Victoria, Western Australia and
South Queensland, and now contains individuals from over 800 colonies. Through
the courtesy of the authorities of the South Australian Museum I have been able
to examine a number of co-type specimens which have been of the greatest assistance
in clearing up many doubtful identifications. In addition to these, I have been
‘able to study many species from localities not represented in my own collection.

For various reasons it has not been found practicable to study the species contained
in these collections in systematic order; in many instances imagines, which are
sometimes essential to satisfactory determination of species, are wanting; in others,
reference to types or authenticated specimens of previously described species is
necessary in order to obviate the possibility of creating synonymy. It is proposed,
therefore, to deal with the various species more or less at random, leaving the pre-
paration of a more comprehensive work until such time as our knowledge of the
Australian termite fauna is much more complete than it is now.

Dr. Eric Mjéberg (1920) has recently added no less than 36 new species to
those previously known from Australia and New Zealand, and has otherwise much
increased our knowledge of the order. He lists 80 species, exclusive of some which
have been too imperfectly described for identification, as comprising the termite fauna
of Australia, Tasmania and New Zealand. To this list, however, should be added
two old described species, v7z., Calotermes convexus, Walker, and C. obscurus, Walker,
the types of which are still in existence, Leucotermes paradoxus, Frogg., and eight
Northern Territory species described by me in 1915.

Eight new species, one new variety, and one hitherto unknown imago are described
in the present paper, leaving at least 20 new species in the writer’s collection to be
dealt with. In addition to the latter number there are several species which cannot
be satisfactorily described until more complete material is available for study.

A critical examination of a large collection of Hamitermes, Drepanotermes, Leuco-
termes, Rhinotermes and Coptotermes, has shown conclusively that in many species
determinations cannot be made satisfactorily, if at all, from the soldier and worker
castes alone. This fact will be made clear in this and subsequent papers.

Although aware of Fuller’s recent paper (1920) on the antennae of termites, I
have followed recent writers on the Australian species in regarding the number and
form of the joints of this organ as possessing considerable taxonomic value. While
due allowance must be made for the variations which certainly do exist in the
antennae of nest-series, and not infrequently in the antennae of individuals, I see
no reason at present for departing from the general practice of referring in some
detail to antennal characters in descriptions of new species.

The wing measurements recorded in this paper were taken from the humeral
suture to the apex. Head lengths, excepting where otherwise stated, are from the
posterior margin of the head to the apex of the mandibles. Measurements are given
in millimetres.

The type series, unless the contrary is stated, are in the author’s collection.

(5296) Wt. P.8/170 1,000 1/22 Harrow G. 75/1. 2E

364 GERALD F, HILL.

Drepanotermes silvestrii, sp. n.

Imago.—Head very dark brown, nearly black; postclypeus, thoracic, and
abdominal tergites Brussels brown; lower surface chiefly antimony yellow; mid
and hind tibiae, pleurae, and lateral blotches on abdominal sternites dark ; wings
dark brown.

Big. 0. Drepanotermes
stlvestvit, sp. n., head and
prothorax of imago.

Head (fig. 1) rounded behind, flat on the summit, glabrous, moderately setose,
Labrum moderately large, swollen on the sides, rounded in front. Anteclypeus,
membranous, pointed in front. Postclypeus rather larger than in D. rubriceps,
Frogg., twice as wide as long. Eyes small, nearly circular (0-329 x 0-376), slightly
projecting. Ocelli large, oval, oblique, well separated from the eyes, anterior margin
in line with anterior margin of eyes. A small deep impression on either side between
the ocelli and clypeus. Fontanelle large, broadly oval, very distinctly visible, with
small indistinct forward extension. Antennae 18-jointed, not variable in size and
shape of segments ; Ist joint long, stout, cylindrical, more than twice as long as
wide ; 2nd three-fifths the length and two-thirds the width of Ist; 3rd and 4th
very short, shortest of all, narrower than 2nd, coalesced ; 5th short but distinctly
longer and wider than 3rd and 4th; 6th longer and wider than 5th; 7th, 8th and
9th equal ; 10th longer than 9th; 11th to 15th about equal; 16th and 17th a little
longer and narrower, 17th narrower than 11th to 16th; 18th as wide as, but longer
than 17th, as long as Ist, pointed.

Fig. 2. Drepanotermes silvestyii, sp. n., hind
tarsus of imago.

Prothorax (fig. 1) nearly flat, slightly sinuate and bent up in front, antero-lateral
angles rounded, sides rounded to the rounded posterior margin, moderately setose.
Meso- and metathorax uniform brown, wing-stumps similar to those of D. rubriceps
but smaller. Legs (fig. 2) very long and slender, armature alike in each excepting
that the first tibiae bear the usual additional spur ; fourth tarsal of each leg very long
and slender ; femora all about equally stout.

ON SOME AUSTRALIAN TERMITES. 365

Wings (fig. 3) with the margin ciliate, membrane with very few hairs; dark
brown, a little lighter on costal border; subcosta very short, hardly extending
beyond suture ; costa and radius well separated, the latter very dark, joining the
former near the apex, sometimes giving off beyond the middle numerous small veinlets
to the costa ; median of the fore-wing branching from the radius within the wing-
stump, moderately thick at the base, nearer to the cubitus than to the radius, branching
very irregularly, sometimes dividing into two before the middle and each branching
again into two or three, the main superior branch joining the radius near the distal
end of the latter or bending downwards and joining the wing margin at or very
near the apex, if the former, a number of small cells are formed beyond the junction ;
sometimes the branches are all curved downwards and reach the hind margin below
the apex. There is always a network of veinlets between the median and radius.
The median vein of hind-wing branches from the radius beyond the suture, but
otherwise it is like that of the fore-wing; sometimes there is an inferior branch
near the base which joins the cubitus about the proximal third of the wing ; some-
times this branch joins the second one and forms a large elongate cell, or there may
be a succession of cells of varying shape and size. The cubitus of the fore-wing
has from 10 to 16 branches, forked or simple, all of which join the hind margin before
the distal fourth or fifth of the wing: that of the hind wing has from seven to nine
simple or forked branches, the first five to seven of which are much darker than
the others.

Fig. 3. Dyrepanotermes silvestrii, sp. n., wings of imago.

Abdomen large, distended, with eight distinct broad dark tergites, clothed with
short fine hairs, and with apical and lateral margins fringed with longer yellow
hairs. Ventral surface of female with six visible yellow-ochre sternites, each with
dark lateral blotches, the fifth longer than the preceding ones, but much shorter than
the sixth. The male has eight distinct sternites, the fifth and sixth longest, the
seventh much shorter and narrower, the eighth shortest and very narrow. Cerci
short and stout. Styli absent in both sexes.

Measurements* : Length with wings, 19-20; length without wings, 11; head,
with mandibles, 2-16 long; head, at and including eyes, 1-7 wide; antennae, 3;
mandibles, right,0-92-1-03 long, 0-7—0-75 wide; left, 1 -03—1-08 long, 0-62-0-65 wide ;
prothorax, 0-94 long, 1-64-1-78 wide ; forewings, 15-16 long, 4 wide; hindwings,
14-5 long, 4:5 wide ; tibia (i) 1-42, (ii) 1-55, (iii) 2-11 ; abdomen, 2-75 wide.

* Given in millimetres.
5296 2E

366 GERALD F. HILL.

Queen.—Antennae generally complete, 7.e., 18-jointed ; wing-stumps often muti-
lated ; two or three legs generally amputated, claws of remaining legs short and
blunt, apices of second and third tarsi heavily chitinised. Eight dorsal and six ventral
abdominal plates distinct; pleurae and integument cream-coloured. Abdomen
16 long by 6 wide. Other characters as in imago.

King.—Uniformly dark above; abdomen much contracted and plates over
tapping ; pleurae obscured by dorsal and ventral plates. Other characters as in
queen and imago. One specimen examined ; apparently very old; associated with
two old neoteinic queens.

Neoteinic Queen.—Head and clypeus yellow-ochre, prothorax a little lighter ;
tergites and sternites amber-yellow, the latter with wearing surfaces dark and
heavily chitinised, pleurae and spaces between plates cream-coloured. Head wide ;
eyes small, hardly projecting, only inner facets pigmented; ocelli as in imago ;
fontanelle a large oval cavity about as large as and shaped like eye (in nymphs of the
first form it is much smaller and proportionately more elongate, asin adult) ; antennae
17-jointed, apparently never mutilated, third and fourth joints fused, short ; pro-
thorax shaped as in imago; wing-buds long, slender, subequal, three-fifths length
of the entire sclerite. Abdomen with cuticle horizontally striate.

Measurements: Head, at and including eyes, 1-70 wide; prothorax, 0-94
long, 1-60 wide.

Described from two old individuals found in association with the true king des-
cribed on a preceding page. Several other similar specimens seen. These forms are
derived from nymphs of the second form, 7.e., in the stage preceding the acquisition
of the long wing-buds characterising nymphs of the first form, 7.e., those which are
destined to develop into winged imagines. In second form nymphs, and neoteinic
queens developed from them, the mesonotum and metanotum, including wing-buds,
measure 2:35 in length; in nymphs of the first form, 7.e., potential winged
imagines and true kings and queens, the length is 4:25. The antenna in each is
17-jointed.

Fig. 4. Drepanotermes silvestrii, sp. n.,
head of soldier.

Soldier.—Head orange-rufous to Sandford’s brown ; mandibles mahogany-red ;
labrum yellowish, apex hyaline; antennae nearly as dark as head; junction of
segments hyaline ; pro-, meso- and metathorax russet to argus brown; legs and
abdominal tergites light clay-colour.

ON SOME AUSTRALIAN TERMITES. 367

Head (fig. 4) very large and broad, rounded behind, with a pale median suture
from posterior margin forwards, widest behind the middle, sloping in towards the
antennae, bearing a few moderately long reddish hairs, variable in size. Mandibles
very long, falciform, each with a large angular tooth before the middle (fig. 5).
Labrum large, wide at the base, sloping on the sides to the bluntly pointed apex.
Clypeus three times as wide as long, divided into two lobes by a deep and wide mgdian
cleft, which extends posteriorly into the front of the head. Antennae 17- or
18-jointed, generally segmented as in D. rubriceps, Frogg., sometimes third and
fourth joints closely fused and together only equal to sixth in length.

Prothorax (fig. 4) much narrower than head, anterior half rounded and bent up
in front, slightly emarginate in middle, postero-lateral angles rounded, hind margin
rounded and slightly emarginate in middle; the margin clothed with short stout
reddish hairs. Mesothorax narrower and shorter than prothorax, hind margin with
a few stout reddish hairs. Metathorax as wide as prothorax and clothed like meso-

thorax. Legs very long and slender, with scattered reddish hairs, fourth tarsal very
long. Tibial spurs 3: 2:2, asin other Drepanotermes and Hamitermes.

Abdomen elongate, narrow, with scattered stout reddish hairs and a few siender
golden ones on tergites and sternites. Cerci long and slender. Styli present or absent.

Fig. 5. Dyrepanotermes silvestrii, sp. n.,
base of jaws, labrum and clypeus of
soldier.

Measurements: Total length, 6:25-7; head and mandibles, 2-75-3-1 long ;
thorax and abdomen, 4-4-5 long; mandibles, 1-31-1-64 long; head, 1-6 wide ;
1:25-1:35 deep; antennae, 3-29-3-61; prothorax, 0:51-0:56 long, 1-03-1-12
wide ; tibia (i) 1-5, (ii) 1-5, (iii) 2-11-2-39 ; abdomen, 1-5 wide.

Worker.—Colour of head as in soldier, or a little darker, with pale median suture
extending forwards from the posterior margin, widening behind the fontanelle and
spreading out behind the frons ; clypeus clay-colour, labrum yellow-ochre ; antennae
a little paler ; rest of insect clay-colour ; prothorax a little darker than tergites and
legs. Immature workers and soldiers have body and legs tinged with rose pink.

Head large, rounded on the sides and behind, nearly as wide as long, widest behind
the base of the mandibles. Labrum large, convex, swollen on the sides, rounded in
front. Anteclypeus membranous, short, pointed. Postclypeus large, convex at base,
not quite as long as wide, sides rounded, anterior margin truncate. Antennae
very long, 18-jointed, arising within a deep cleft situated well in from sides of head.
Fontanelle as in imago.

Prothorax as in soldier, more setose, much narrower than head. Legs long and
slender, clothed with scattered reddish hairs. Tibial spurs 3: 2: 2.

Abdomen elongate oval, hairs more numerous and more slender than in soldier.
Cerci long and slender.

Measurements: Total length, 7-7-5; head, with mandibles, 2-16-2-35 long ;
thorax and abdomen, 5-5 long; head, 1-93 wide ; antennae, 3-61 ; prothorax, 0-56-
0-6 long, 1-22 wide ; mandibles, left, 0-97 long, 0-65 wide, right, 0-86 long, 0-77 wide.

368 GERALD F. HILL.

Biology.

This is one of the three predominant species of mound-building termites that are
found in the Townsville district, N. Queensland. It inhabits the same localities as
H. perplexus, sp. n., and the termitaria of the two species are often found in close
proximity, although those of the latter are rather more common, especially on _hill-
sides. On the higher and stony localities many large colonies live entirely in under-
ground galleries, the extent and nature of which have not yet been investigated
sufficiently to determine whether they are connected with large masses of cells and
passages comparable with termitaria. This seems most probable, since it is known
that these colonies collect and store considerable quantities of food, and that eggs,
young larvae and the reproductive forms are not found in the galleries near the
surface, which serve apparently only for the accommodation of workers and soldiers,
and a few adolescents of these castes, and for the temporary storage of food material
during and just after harvesting operations. As all the normal castes are reared by
these colonies, it seems reasonable to assume that each is provided with an under-
ground system suitable for the location of the royal pair and their young and for the
storage of food. On the other hand, considerable excavating failed to disclose a
regular nest or “‘ nursery ’”’ in the closely allied species D. septentrionalis, sp.n., in the
Northern Territory (Hill, 1915). In the case of D. septentrionalis, small foraging
parties of soldiers and workers are commonly found in the termitaria of Coptotermes
and Eutermes, but this is not the case with D. silvestriit. Access to the surface is
gained by means of numerous small oval openings, from 18in. to 3ft. apart, and
extending over an area of from 6 ft. to 12 ft. in diameter. These openings measure
about 3 mm. long by 6 mm., and except when actually in use, 7.e., at harvesting or
‘““swarming’”’ periods, are sealed with earthy matter, either level with the surface
or just below it, in either case rendering their detection very difficult. Similar surface
openings are found in the vicinity of termitaria, when these are constructed; but
they appear to be used solely at harvest time—certainly not to provide a means of
exit for the winged forms at the time of swarming. The natural dispersal of the imagines
has not been observed, and it is not known whether the phenomenon occurs during
daylight or at night. A day or two before swarming takes place, slits are cut in the
walls of the termitarium, generally in several places near the outer margin and several
inches above ground level. These slits are sealed over by a projecting crust of moist
earthy matter, as in H. perplexus, sp. n., and remain thus until weather conditions
are favourable for the flight, after which they are cemented up flush with the general
surface of the walls.

One of the most remarkable habits observed in this species, and also in D. septen-
trionalis, is that of gathering food supplies by day as well as by night. Froggatt
(1915) observes that travellers in the bush, who have gathered a mass of dried grass
upon which to make their temporary bed, have been aroused to find hordes of
termites (species not stated) cutting the material into lengths and removing it for
food. Such is by no means a rare experience in North Australia, and it occurs
during the day as well as at night, D. septentrionalis being the species concerned in
all cases which have come under my notice. In this district I have frequently seen
countless thousands of soldiers and workers of D. silvestrii issuing from several
holes in the surface and spreading out in irregular columns over an area of several
yards, each worker cutting off a length of grass (leaf or stem), a piece of eucalyptus
leaf or twig, or seizing a seed or small piece of bark and hurrying back along the
column to one of the openings, at each of which there is a good deal of congestion,
but no sign of disorder or wasted effort. Throughout these operations the soldiers
are much in evidence, regulating the traffic, scouting on the outskirts of the working
parties, attacking marauding ants or any other insects or spiders they may encounter,
and generally taking a strenuous and important part in the proceedings. Their
behaviour is in marked contrast to that of their near allies the Hamitermes, and

ON SOME AUSTRALIAN TERMITES. 369

many other species, in which the soldiers appear to be the embodiment of cowardice
and uselessness. On one occasion harvesting operations were observed at night on
the roadside in one of the more populous residential areas of the town, the material
gathered being almost entirely coarse dry grass, which was cut into pieces about
half an inch in length. The nature of the food varies according to the season of the
year, the flora in the immediate vicinity and other circumstances. When there is
a plentiful supply of dry grass this material appears to be most favoured, but there
is nearly always present a quantity of grass and other seeds and a good deal of
vegetable debris. In the vicinity of eucalyptus trees they gather pieces of leaf, leaf-
stems, twigs and bark. The latter are carried into the termitaria in the rough state
and afterwards dressed into pellets of varying size and shape. None of the species
of this genus are wood-eaters, nor are they known to attack cultivated cereals.

The termitaria are nearly always low, flat and more or less circular masses composed
of intensely hard cement-like material, varying in toughness according to the soil
in which they are situated. In size they vary from about 8 in. to 2 ft. in height
by 2 ft. 6in. to 8 or 9 ft. in diameter. In gross appearance they resemble a mass of
soft mud which has spread over the surface and hardened by evaporation (PI. ix, fig. 1).
There is no well-defined outer casing or wall (Pl. ix, fig. 2) as in nests of Coptotermes,
the whole of the superstructure being composed of similar material. The interior
is occupied by very large flattened chambers connected with each other by small
circular holes large enough to permit of the free passage of soldiers and workers from
chamber to chamber. Similar but rather larger chambers extend below ground,
under the middle of the superstructure, to a depth roughly corresponding to the height
of the latter. These chambers are excavated in the soil, and are much less resistant
to digging operations than those above ground. Below them are several passages
extending more or less vertically into the soil beneath. The majority of the chambers
in the superstructure are occupied by workers, soldiers, and older adolescents, and
by masses of grass and other foodstuffs. The latter is generally stored in the rather
smaller outer cells, many groups of which are reserved for the reception of the waste
matter from the community, ?.e., alimentary rejectamenta and the heads of dead
soldiers. Evidently much of the waste material is carried in the jaws to these
chambers, where it is tightly packed until the space is entirely filled, then the small
entrances are cemented up, apparently never to be reopened. Other chambers are
reserved for the reception of the faecal matter of certain individuals, probably
soldiers and workers, who evacuate directly into them. Such chambers, when in
use, are indicated by a deposit of more or less liquid matter just within the small
entrance hole, the remainder being empty. As the deposit increases and hardens
the entrance becomes blocked and is then cemented up. Analyses of the rejecta-
menta in these termitaria show that they contain about 32 per cent. of inorganic
matter. Additions to these nests are nearly always made by extending the
outer walls without increasing the height, and the increase in the diameter of the
superstructure is greatly in advance of that of the underground portion. The latter
is occupied by the reproductive forms, eggs, larvae and nymphs, and by their
attendant soldiers and workers.

All the different castes have been found in the same nest and at the same time,
but an ovigerous neoteinic queen has not been found in a nest presided over by a
true queen. The soldiers are very numerous, active and pugnacious, and are capable
of a most effective fight against marauding ants. When the termitarium is broken
into the soldiers rush out in all directions, attacking every animate object they come
in contact with—their fellow-soldiers and workers, ants, lizards and one’s hands
Teceiving equal attention. Some few devote their energies to rescuing their defence-
less larvae, but this function devolves more upon the workers, who are hardly less
pugnacious. Normally the colony is presided over by one true queen, who is generally
located at, or just below, ground-level in a flattened cell of rather smaller size than
the average cell in these nests. In one case only has a true king been found in the

370 GERALD F. HILL.

queen cell. The true queen produces an enormous number of eggs, which are carried
away by the workers and stored in masses in cells near the walls or near the queen-
cell. Egg-laying is not confined to one particular season of the year, but it is not a
continuous process, since eggs are often absent in certain thriving colonies while
present in great numbers in others close by. Neoteinic queens are substituted for
a true queen when a colony is naturally or designedly orphaned. In one colony
a true king was found in a large cell with one ovigerous neoteinic queen ; in another
there were one true king and two of these neoteinics, in a third there were two neoteinics
only, and in a fourth one neoteinic king and two neoteinic queens. Neoteinic queens
of this species produce neoteinic males and females, as well as soldiers, workers, and
nymphs of the first and second form.

The following field notes refer to termitaria of this species which have been kept
under observation for some time :—

(1) This colony was orphaned on 22nd August 1919. When examined on 15th
June 1920, it contained 20 young neoteinics of both sexes. There were no eggs or
very young larvae present, but there were numerous half-grown larvae and second form
nymphs. On 26th October 1920, two ovigerous neoteinic queens and four neoteinic
males were removed from the nest. Eggs, young larvae and nymphs of the second
form were plentiful. Some of the latter and one young neoteinic female were left
in the nest, with workers and soldiers. By 15th February 1921, the termitarium
was again restored to its original size and presented a very prosperous appearance.
Eggs and young larvae were present, but no gravid female could be found. There
were no young neoteinics present, but the second form nymphs which were left in the
nest on the 26th October had now developed into nymphs of the first form. The
parent of the eggs and young larvae found on this date was presumed to be the young
neoteinic female left in the nest on 26th October.

(2) This nest was orphaned on 15th June 1920. On 26th October 1920, it con-
tained four ovigerous neoteinics and nine immature neoteinic males and females.
There were present also numerous second form nymphs, besides the usual workers and
soldiers. The termitarium was now completely destroyed. On 15th February 1921,
the nest was found to have been rebuilt to its original size and to contain numerous
eggs, young larvae and nymphs of the second form. There were no nymphs of the
first form or imagines present and the parent of the eggs and young larvae was not
found. The whole termitarium was again destroyed, and on 18th March 1921
a good deal of it was found to have been rebuilt.

Each of several other nests which were orphaned at different periods of the year
were found to contain neoteinic queens when examined subsequently. It has not
been ascertained if a colony once deprived of its true queen is ever again presided
over by another true queen ; the contrary appears to be the case. In nests which are
presided over by a true queen, or by one or more gravid neoteinics with numerous
neoteinics in reserve, nymphs of the second form are found throughout the year, except-
ing from the middle of December to the end of January. A moult takes place about the
former period, and the resulting first form nymphs have been found as late as 13th
February, but the majority undergo their final moult and appear as imagines about
the beginning of January, and all have moulted before the 8th March. When true
queens or mature neoteinic queens are not present, nymphs of the second form may
be present throughout the year. First form nymphs have not been found later than
15th February or earlier than 5th November. The actual date of swarming is
determined by rainfall. In 1919-1920 first form nymphs were plentiful in the nests
on 5th November 1919; the final moult took place between this date and 8th
December, when most of the imagines were capable of flight, although some had not
yet moulted. On 6th January and 15th January (1920) first form nymphs and
imagines were still present in the nests. The former moulted before 30th January,
and swarming took place before 10th February. In 1921 the wings of the majority

ON SOME AUSTRALIAN TERMITES. 371

were fully developed on 15th February, but on 8th March, up to which date only
light rain fell, these forms were still present in all the nests examined. On this
date the tips of the wings showed marked damage due to prolonged occupancy of
the parent nest. Similar conditions prevailed in the nests of Hamitermes perplexus,
sp.n. Heavy showers fell on the night of the 9th March and throughout the day and
night of 10th. Swarming of H. perplexus took place during the afternoon and evening
of the latter date. The swarming of D. silvestrii was not observed, but probably
took place about the same time, since none of the nests contained imagines on
14th March.

It is not intended to discuss in detail here the numerous other forms of life which
have been found in termitaria, but brief mention may be made of one species which
plays an important part in the economy of two species of termites dealt with
in this paper. At a very rough estimate it may be said that 80 per cent. of the
termitaria of D. silvestrit and H. perplexus are invaded and permanently occupied
by the very common and widely distributed ant, Iridomyrmex sanguineus, Forel,
which is particularly abundant on the low-lying country in the vicinity of Townsville.
The termitaria are entered by means of holes burrowed into the walls (PI. xii, fig. 2),
in and out of which pass endless streams of ants in their journeys from one nest to
another. If a termitarium is cut open vertically, it will be found that the ants
have greatly enlarged the original galleries so as to form large flattened chambers
in tier upon tier, until finally the greater part of the structure is in their undisputed
possession. The floor of each cell is thickly covered with the eggs, larvae and
pupae of the invaders, and immense numbers of ants throng all parts not actually
in possession of the termites. As the ants extend their sphere, the termites are driven
back from chamber to chamber and destroyed, until but a few stragglers are left.
The complete, or nearly complete, occupation of a termitarium is evidently a matter
of time, during which the advance is being constantly delayed by the termites walling
up their galleries and passages as they retreat. The remains of the dead termites
in the chambers occupied by ants show clearly that the nests are not attacked merely
to provide a dry and safe shelter, but that the original occupants are used as food.
Immediately the walls are broken with the pick the ants swarm out in countless
thousands, destroying and carrying off the dislodged termites, crawling up one’s
legs and attacking one’s hands, head or any skin surface to which they can gain
access. Others of their kind gather from all directions to take part in the onslaught,
until the nest and the surrounding ground is a seething mass of insect life. Under
these conditions a close examination of the nest or its occupants is impossible, and
it is only by finding an ant-free nest that one can hope to investigate its interior.
Within a few minutes of the nest being broken into all the neighbouring ant-
infested termitaria of these two species will be found to contain the bodies of freshly
killed termites, while files of ants pass to and fro so long as a termite remains exposed
to attack. Plate xii, fig. 1, shows ant tracks made on the surface of the ground
approaching a mound of H. perplexus. It is a remarkable fact that the mounds
of a certain species of Eutermes, which are very common amongst those of the
Drepanotermes and Hamitermes, are never molested by Iridomyrmex.

The imagines of the beetle, Cryptodus grossipes, Fairm., have been found in the
cells of D. silvestrit, and Mandalotus germinatus, Lea, has been taken on two occasions
in the nests of H. perplexus. Their relationship to their hosts isnot known. Bubaris
indemnis, Pascoe, has been found under the walls of termitaria of several kinds,
but they appear not to come directly into contact with the termites.

Ciliates (? Trichonympha), which occur in vast numbers in all the workers
and soldiers and in many imagines of Mastotermes davwinicnsis, Frogg., in
Townsville, have not been found in Drepanotermes silvestrit, or in any other
locai species of termite.

372 GERALD F. HILL.

Drepanotermes septentrionalis, sp. n.
Termes rubriceps, Hill (nec Frogg.), Proc. Linn. Soc. N.S.W., xl, pt. 1, 1915.

Imago.—Head bay, clypeus argus brown; labrum, palpi, antennae and legs
buckthorn brown, anteclypeus lighter; thorax and abdominal tergites auburn ;
lower surface of abdomen uniform ochraceous tawny ; wings Brussels brown, faintly
tinged with yellow behind second vein. ‘

Head wide, rounded behind and on the sides, flat on the summit, moderately
hairy. Labrum moderately large, swollen on the sides, rounded in front. Ante-
clypeus yellow, membranous, slightly pointed in front. Postclypeus convex, twice
as wide as long, with median suture very distinct. Eyes small, circular (0-376 dia-
meter), prominent. Ocelli broadly oval, widely separated from eyes. The small
deep impression between ocelli and clypeus pale-coloured. Fontanelle broadly oval,
about the size and shape of ocelli, with short indistinct forward extension, similar
to that of D. silvestrii, sp. n., but slightly larger. Antennae 18-jointed ; Ist segment
moderately long and wide ; 2nd half as long as Ist ; 3rd, 4th and 5th small and closely
fused; 3rd and 4th equal to each other, and a little longer than 5th; 6th longer
and wider than Sth. ,

Thorax similar to that of D. silvestri1, but more rounded on the sides, similarly
clothed. Legs as in D. silvestrit.

Fig. 6. Dyvepanotermes septentrionalis, sp. 0.,
wings of imago.

Wings (fig. 6) with the margin ciliate; membrane with many hairs, subcosta
very short, hardly extending beyond suture, costa and radius well separated, the
latter very dark and connected with the former near the apex of the wing by a few
indistinct nervures ; median of the fore-wing branching from the radius within the
wing-stuimp, that of the hind-wing just beyond suture, nearer to cubitus than to radius ;
branches of the median and cubitus very irregular and not alike in either fore- or
hind-wing.

Abdomen elongate, nearly cylindrical, moderately densely clothed with short
reddish hairs; ten dorsal and six ventral plates distinctly visible. Cerci as in
D. silvestrit.

Measurements: Length with wings, 15; length without wings, 8; head, with
mandibles, 2-06 long ; head, at and including eyes, 1-78 wide ; prothorax, 1-03 long,
1-22 wide; fore-wings, 13 long, 3-25 wide; hind-wings, 12-5 long, 3-5 wide ;
abdomen, 1-73-1-92 wide.

Soldier.—Very like D. silvestrii, sp. n., from which it differs in having the labrum
larger and more rounded at the apex, clypeus shorter and less strongly lobed, antennae
(fig. 7) of the same number of joints, 7.e., 17 or 18, but the size and shape of the basal
joints very distinctly different, viz., in D. septentrionalis the 1st joint is shorter and
wider, the 2nd shorter and narrower, the 3rd and 4th very short, together equal

ON SOME AUSTRALIAN TERMITES. 373

to the 3rd in D. silvestrit, 5th about half the length of corresponding joint in
the latter species. The head very slightly redder than in the allied species. In
size it is intermediate between individuals of D. silvestrit from high stony localities
(Castle Hill, Townsville), and those from the low-lying country in the vicinity
(Townsville Common).

Fig. 7. Dvrepanotermes septentrionalis, sp. n.,
basal joints of antenna of soldier.

Worker.—Very like that of D. silvestrit; head more reddish, median suture very
obscure, not widening in front to surround the fontanelle ; fontanelle hardly visible ;
antennae with 18 joints, stouter, but otherwise similar.

This caste, like the soldier, is intermediate in size between D. silvestri1 from the
hill-sides and from the plains, as shown by a series of measurements of antennae,
mandibles and tibiae.

Biology.
he imago was originally described under the name of Termes rubriceps, Frogg.,
from a de-alated female taken on 11th January 1914 (Hill, 1915), and is here re-
described from a perfect specimen of the same sex taken under similar circumstances
and in the same locality on 3rd February 1918.

From a very thorough knowledge of all the country on either side of the Darwin-
Katherine Railway within 60 miles of the coast, Iam convinced that these termites do
not construct termitaria, but live in rambling underground galleries as previously
described (Hill, 1915).

Drepanotermes daliensis, sp. n.
Soldier.—Head very dark, nearly black ; front of head, clypeus, anterior part of

prothorax and mandibles a little lighter, clypeus yellow; antennae, palpi and legs
ochraceous tawny.

Fig. 8. Head of soldier of (a) Dvrepanotermes

fo)
daliensis, sp. n.; (b) D. perniger, Frogg.

Head (fig. 8,a) very large, widest behind, sloping in slightly to the base of the jaws ;
frons flattened, a little rugose, median suture indistinct. Mandibles very long and
slender, falciform, each with a large angular tooth nearer to the base than to the apex

374 GERALD FF.) HIEL:

and generally directed slightly forward. Labrum large, convex, rounded on the sides
to the bluntly rounded apex. Clypeus large, slightiy convex, emarginate in front,
divided medially by a deep depression. Fontanelle very indistinct. Antennae
(fig. 9,b) very long and slender, 19- or 20-jointed, rarely 18; Ist joint twice as long
and about half as wide as 20d; 2nd nearly cylindrical, one-third longer than
3rd; 3rd narrowest at base, wider than 4th at apex; 4th shortest of all; Stha
little longer and wider than 4th; 6th neariy as long as 2nd; 7th longer and
narrower than 6th.

Fig. 9. Proximal segments of antenna of soldier of
(a) Drepanotermes perniger, Frogg.; (b) D. daliensis,
sp. n.

Prothorax similar to that of D. silvestrii, but anterior half narrower. Legs long
and slender, with scattered hairs except on inner side of tibiae, which are fringed with
longer and stouter hairs ; mid-tibiae with two short stout apical setae on upper side,
which are absent in hind tibiae; first three tarsals very short, 4th very long ;
tibial spurs 3: 2: 2.

Abdomen as in D. silvestrit, sp. n.

Measurements :—
D. daliensis, sp. n. D. perniger, Frogg.

Head and mandibles, long 3- 290-3 - 807 3-196
Thorax and abdomen, long 3-290 3-290
Mandibles, long 1-551 1-598
Head, deep 1-475 1-034
» wide 1-645 1-598
Antennae 3-666 3-666
Prothorax, long : 0-658 0-564
a wide 1-128 1-081
Tibia (i) 1-410 1-457
imi ae) 1-457 1-363
elit) 2-115 - 2-303

Abdomen 1-05 ae =

Worker.—Colour of head ¢s in soldier ; clypeus and jaws (excepting teeth) tawny
olive ; labrum, palpi, antennae, thorax and legs clay-colour.

Head large, rounded behind and on the sides, widest near the middle, a pale coloured
and very distinct median suture extending from back of head forwards, spreading
out around the fontanelle, which is a sharply defined, small, broadly oval depression ;
frons sloping slightly to the base of the clypeus, very faintly rugose. Labrum large,
very convex, covering apex of jaws, narrower than anteclypeus at base, swelling out
sharply in the middle to the rounded apex. Anteclypeus large, middle of anterior
margin produced into a point. Postclypeus large, twice as wide as long, convex,
truncate in front, strongly arcuate behind, median suture hardly visible, with
very few hairs. Antennae 19-or 20-jointed; 3rd, 4th and Sth joints short ;
4th shortest.

Prothorax as in soldier, entire surface with stout reddish hairs. Legs long and
slender, as in D. silvestrit, sp. n.

ON SOME AUSTRALIAN TERMITES. 375

Abdomen narrow, tapered to the pointed apex, with pale reddish hairs of various
lengths. Cerci long and slender.

Measurements :—
D, daliensts, sp.n. D. perniger, Frogg.
Total length “ag ar 6
Head, long Ay a 2°115—2-'250 ae 1-927
Thorax and abdomen, long 3:995 rs 3°920
Head, wide WA ca 1-739 fe 1-692
Antennae .. aes art 3:700 i 3-800
Mandibles :
left ae aie a4 -987 long by of -890 long by
‘658 wide -640 wide
right 4s »: ok -893 long by wi -799 long by
-752 wide -750 wide
Prothorax, long .. a, 0-705 0-611
Ae wide .. ” 1-034 1-128
Abia o()! «F< io ses 1-316 1-363
is 6) meee si ie 1222 1-316
oe Te (ite Bie ahs : L924 2-068

This species is very closely SaNies to D. permger, Frogg. The soldiers are dis-
tinguished as follows: In D. daliensis the head is very much darker ; the frons is
slightly protuberant and rugose, the middle falling gently into the frontal opening.
In D. perniger (fig. 8,b) the frons is only slightly protuberant, but more rugose. The
clypeus in D. dalzensis is less lobed in front and the furrow dividing it medially is
narrower and shallower; the labrum is much shorter and rounder; the antenna
has always one, but generally two or three, additional joints, the fourth and fifth
of which are very much shorter than in D. perniger (fig. 9,2). The worker may be
distinguished from that of D. perniger by its darker head and 19- or 20-jointed
antennae.

Type series in South Australian Museum, co-types in author’s collection.
NORTHERN TERRITORY: Upper Daly River (H. Wesselman).

Drepanotermes perniger, Frogg.

In his discussion of this species Dr. Mjéberg (1920, p. 69) remarks that the soldiers
from different localities show considerable differences in the colour, size and shape of
the head, but that he has found no constant characters which justify him in regarding
the pale-headed forms from North Queensland and Kimberley as specifically distinct
from the typical dark-headed forms. Then follows a description of the imago, but
unfortunately no locality is given, nor is it stated if his specimens were associated
with pale- or with dark-headed soldiers. On page 57 the same author gives a key for
the differentiation of the soldiers of the two hitherto described species of Drepanotermes,
viz., D. permger, Frogg., and D. rubriceps, Frogg., the former being distinguished
by the very long jaws and very broad, projecting tooth, and the latter by shorter jaws
and triangular tooth. The jaws of D. perniger are figured on page 76.

In this paper I have referred to the similarity which exists in the heads of soldiers
of certain species the imagines of which show marked specific differences ; for this
reason I cannot agree with the suggestion that pale- and dark-headed forms are
referable to a single species. If the imagines described by Dr. Mjéberg as D. perniger
were associated with pale-headed soldiers it is most probable that they are referable
to another species. With regard to characters given for the soldiers of D. perniger
and D. rubriceps, it may be said that the shape of the mandibular tooth is
variable in specimens from the same colony, and that the form figured by
Dr. Mjéberg for D. perniger is one commonly found in D. rubriceps, D. silvestrii,
D. septentrionalis and occasionally in D. daliensis.

376 GERALD F. HILL.

Specimens of soldiers and workers in the South Australian Museum from Beverley,
W. A., Moorella, C.A., and Leigh Creek, C.A., agree perfectly with co-types of
D. perniger, Frogg., and are undoubtedly referable to that species. I have not
had an opportunity of examining specimens of Drepanotermes from Kimberley, nor
dark-headed forms of the genus from Queensland.

Drepanotermes rubriceps, Frogg.
Termes rubriceps, Froggatt, Proc. Linn. Soc., N.S.W, xxii, 1897, p. 730.

In an earlier paper (Hill, 1915) I described a de-alated imago under the above name,
the determination of the species having been made for me by Mr. Froggatt, from
soldiers and workers only. Later, a perfect specimen was secured from the same
locality (Darwin, Northern Territory) and under similar conditions. Recently I
have had for examination a series of imagines, soldiers and workers from Tennant’s
Creek and Leigh Creek, Central Australia (South Australian Museum Collection),
which, although not compared with the type soldiers and workers, I consider to be
D. rubriceps, Frogg., the type locality of which is McKinley Ranges, Central Australia.
A comparison of the winged forms from Darwin with those from Central Australia
shows that they are referable to two quite distinct species. The structural differ-
ences between the respective soldiers, however, are hardly appreciable. The heads of
the Central Australian specimens are distinctly lighter than those of the Northern
species and there are small differences in the structure of the antennae.

Fig. 10. Drepanotermes
vubriceps, Frogg., head
of imago.

The following is a description of the imago of this species. The Northern species is
described in the preceding pages under D. septentrionalts.

Imago.—Head very dark brown ; clypeus yellow-ochre ; antennae darker than
clypeus, brownish ; trophi and ventral surface antimony yellow ; prothorax yellow-
ochre, with brown blotches; wing-stumps and abdominal tergites argus brown,
the former with dark vein bases ; wings brown, radius and branches of the cubitus
very dark, costal margin pale.

Head (fig. 10) large, rounded behind, flat on the summit, clothed with numerous
short, fine, pale hairs. Labrum narrow at base, swollen on the sides, rounded in
front. Anteclypeus membranous, pointed in front. Postclypeus moderately large,
twice as wide as long, convex, rounded behind, truncate in front, sloping on the sides.
Fontanelle very small, lanceolate, with anterior end bifurcated. Eyes large, pro-
jecting moderately, nearly circular (0-610 x 0-470). Ocelli large, oval, oblique,
a little less in length than short diameter (horizontal) of eyes, very close to the eyes,
their anterior margin a little posterior to the anterior margin of eyes. Mandibles with
dentition as in D. silvesirii, sp. n. Antennae very long and slender, 16-jointed ;

ON SOME AUSTRALIAN TERMITES. OV My

Ist joint more than twice as long as 2nd; 3rd about as long as 2nd, narrower and
more turbinate ; 4th a little shorter than 3rd, wider, oval, always shortest of all, but
rarely markedly so ; 5th longer and wider than 4th ; 6th a little shorter and narrower
than 5th; 7th to 9th, inclusive, increasing gradually in length ; 10th to 14th about
equal to each other, very little longer than 9th; 15th and 16th slightly longer than
14th, equal to each other ; ; joints 5 to 16, inclusive, more or less stalked; 5 to 15
slightly turbinate ; 7 to 16 distinctly slender ; number and shape of joints apparently
very constant.

Prothorax roughly triangular, setose, nearly flat, wider than long, anterior margin
slightly arcuate, bent up in the middle, antero-lateral angles rounded, sides nearly
straight, sloping sharply to the rounded posterior margin, hairy. Mesothorax with
a broad dark stripe down the middle ; wing-stumps large, setose, about two-thirds
as long as the mesonotum. Metanotum similar, but wing-stumps small, half as
long as the visible portion of the metanotum. Legs as in D. stlvestrit (fig. 2).

Wings (fig. 11) very large and broad ; fore-wing a little longer and a little narrower
than hind-wing, margin ciliate. Anterior margin distinctly yellow-ochre. Costa
and radius dark at base, becoming yellowish further on, the latter well separated

Fig. 11. Dyrepanotermes rvubriceps, Frogg., wings of
imago.

from the former in the proximal half of the wing, closer but distinctly separated in
the apical half to near its junction just before the apex. Median dark at the base
only, the rest very indistinct, running much nearer to the cubitus than to the radius.
In the fore-wing the median separates from the radius within the wing-stump, runs
straight to the apex of the wing, giving off five or six ill-defined superior branches.
In the hind-wing the separation from the radius takes place beyond the suture,
there are seven or eight very obscure branches, those near the base being short,
the others running out to the wing margin, which they join just above or below the
apex. The cubitus of the fore-wing joins the hind-margin a little below the apex
and gives off from 12 to 15 branches, the first 9 to 12 of which are very distinct
and some are forked. In the hind-wing there are generally 12 branches, simple or
‘forked once or twice. Other variations are common.

Abdomen large, showing nine distinct tergites, each clothed with fine, short, pale
hairs and a fringe of large golden hairs on posterior margin. Sternites with indis-
tinct brownish patches laterally. Cerci very short and broad.

Measurements: Length with wings, 21-22; length without wings, 11; head,
with mandibles, 2-35 long ; head, at and including eyes, 1-88 wide ; antennae,
3:29-3:75 ; mandibles, right 0-94 long, 0-75 wide; left, 0-94 long, 0:6 wide;
prothorax, | long, 1-78 wide; fore-wing, 18-3 long, 5 wide ; hind-wing, 17-5 long,
9°25 wide ; tibia (i) 1-36, (i i) 1-5, (iii) 2; abdomen, 3-5 wide.

378 GERALD F. HILL.

Soldier.—Head (fig. 12) yellow-ochre to ochraceous orange; labrum whitish
yellow ; clypeus with anterior margin bordered with hyaline membrane ; mandibles
a little darker than head, darkest at tip; remainder of insect pale yellow.

Antennae very long and slender, 17- or 18-jointed; 1st joint long, rather more
than twice as long as 2nd and one-third wider; 3rd as long as 2nd, but narrower ;
4th shorter than 3rd but equally wide, more or less fused with it; 5th as long as
3rd, narrow; 5th to 9th increasing successively, all elongate, narrow; 10th to
18th about equal in length, a little shorter than Ist.

Measurements: Head and mandibles (crossed), 2:82 long; head, 1-73 wide ;
antennae, 3-57; thorax and abdomen, 3-29 long; prothorax, 0-6 long, 1-13 wide ;
tibia (i) 1-27, (ii) 1-45, (iii) 2-06.

The imago differs from D. silvestrit in having a larger body, wings longer and
paler, lower surface of abdomen paler, much smaller and differently shaped fontanelle,
larger eyes, fewer joints in antennae (16 as against 18 in D. silvestri1), segmentation
distinctly different, head and prothorax much lighter coloured. The soldier is
lighter in colour than that of D. silvestrii, and the front of the head a little less rugose ;
otherwise there is little to distinguish them.

Fig. 12. Drepanotermes rubriceps, Frogg.,
head of soldier.

From D. septentrionalis, sp. n., the imago is easily separated by its larger size,
longer and darker wings, darker prothorax, and two less joints in the antennae.
From D. perniger, Frogg., as described by Mjéberg, it differs in being much larger,
having much longer wings, and also in the eyes, fontanelle, and antennae.

Type series of imagines in South Australian Museum; co-types in author’s
collection.

CENTRAL AUSTRALIA: Tennant’s Creek (J. F. Field); Leigh Creek, Everard
Ranges (S. A. White). WESTERN AUSTRALIA: Mullewa.

Hamitermes parvus, sp. n.

Imago.—Clypeus, meso- and metathorax, legs, abdominal sternites and tergites
mummy-brown ; prothorax much darker brown ; mouth-parts clay-coloured ; tarsi
pale stramineous ; claws pale ferruginous ; wings brown.

ON SOME AUSTRALIAN TERMITES. 379

Head (fig. 13) clothed with numerous short and moderately long hairs, longer
than wide, rounded behind and on the sides to the base of the jaws (widest part),
flat on the summit. Fontanelle in the form of two elongate straight clefts in the
middle line. Eyes small, hardly projecting beyond sides of head. Ocelli oval and
widely separated from the eyes. Antennae 14-jointed, arising from a raised tubercle
within a deep and wide fossa in front of the eyes, the tubercle being about equidistant

Fig. 13. Hamitermes parvus, sp. n., head of
imago (a) in profile; (b) from above.

between the anterior margin of the eye and the postero-lateral margin of the clypeus ;
Ist joint twice as long as 2nd, 3rd shortest, indistinctly separated from the 2nd and
Ath. Labrum rather narrow at the base, swollen on the sides, rounded in front.
Anteclypeus arcuate anteriorly, slightly convex, half as long as wide, half as
wide as postclypeus, divided medially by a suture which extends across the latter.
Postclypeus large, convex, anterior margin slightly concave, posterior margip
semi-circular, greatest length slightly more than greatest width.

Prothorax nearly twice as wide as long, very slightly convex and bent up along
the anterior margin, antero-lateral angles rounded and slightly bent up, sides sloping
sharply to the nearly straight posterior margin. Legs with fore tibiae and all femora
stout; tibial spurs 3: 2-2,

Wings (fig. 14) brown, large veins and first five or six branches of the cubitus
well defined ; entire margin excepting proximal fifth of hind margin ciliate; veins
with a few long hairs along the entire length, venation variable and not always
alike in either fore- or hind-wings of the same individual ; cross suture straight.

Fig. 14. Hanvitermes parvus, sp. n., wings of imago.

Abdomen nearly cylindrical; seventh sternite of male much longer and darke
than corresponding sternite of female, densely clothed with short fine reddish hairs.
Cerci short, conical ; basal segment very broad (0-8) and flattened.

Measurements: Length with wings, 6-25-7; length without wings, 3-25-4 ;
head, with mandibles, 0:75-0:84 long ; thorax and abdomen, 2:63 long; head, at

(5296) 2F

380 GERALD F. HILL.

and including eyes, 0:65-0:67 wide; antennae, 1-08; mandibles, right, 0-41 long,
0-28 wide, left, 0-4 long, 0-25 wide ; prothorax, 0-28 long, 0-51 wide ; wings, 5 long,
1-27-1-4 wide; tibia (i) 0-51, (ii) 0-48, (iii) 0-7.

Soldier.—Head buff yellow; antennae, palpi, thorax and legs paler; abdomen
grey (due to stomach contents) ; jaws castaneous, with basal half much paler.

Head (fig. 15) rounded behind and on the sides, compressed dorso-ventrally,
clothed with scattered short reddish hairs. Mandibles short and stout, with a blunt
angular tooth on each about the middle. Labrum broad at the base, sloping in to the
rounded apex. Clypeus broad and rather indistinct. Antennae arising from raised
tubercles close to the base of the jaws, 13-jointed; Ist joint nearly twice as long
as 2nd; 2nd and 4th equal; 3rd half the length of the 2nd and 4th; Sth and 8th
three-fourths the length of the 4th ; 9th-11th equal to the 2nd; 12th rather longer ;
13th longest, twice as long as 2nd and 4th.

Fig. 15. Hamitermes parvus, sp. n., head of soldier
(a) from above ; (8) in profile.

Prothorax similar to that of worker. Legs short and moderately stout ; tibial
spurs 372: 2.

Abdomen elongate-oval, clothed with numerous moderately long and a few very
long reddish hairs.

Measurements: Total length about 3:3; head and mandibles, 1-22 long; thorax
and abdomen, 2 long ; mandibles, 0:47 long ; head, 0-65 wide, 0:47 deep ; antennae,
0-86; prothorax, 0:18long, 0-47 wide; tibia (i) 0-46, (ii) 0-37, (ii1) 0-56; abdomen,
0-7 wide.

Worker.—Head creamy ; mouth-parts, thorax and legs paler; abdomen nearly
hyaline.

Head almost spherical, slightly longer than wide, clothed with scattered red
hairs; labrum large convex, rounded in front ; clypeus similar in shape to that of
winged form, at each end a ferruginous spot (articulation of mandible). Antennae
13-jointed, segmented similarly to that of winged form, arising from a raised tubercle.
Dentition as in imago.

Prothorax nearly twice as wide as long, arcuate and bent up in front. Legs
stout; tibial spurs 3:2: 2.

Abdomen elongate-oval, bluntly rounded at apex, clothed with moderately long and
stout hairs, with a few much longer ones scattered over dorsal and ventral surfaces.
Cerci large and prominent, base with several very long slender hairs.

ON SOME AUSTRALIAN TERMITES. 381

Measurements : Total length (about), 3-47; head with jaws, 0-7 long; thorax
and abdomen, 2-68 long ; head, 0-64 wide, 0:32 deep; antennae, 0:98 ; mandibles,
left, 0-38 long, 0-25 wide, right, 0-36 long, 0-28 wide; prothorax, 0-22 long,
0-4 wide ; tibia (i) 0-46, (ii) 0-38, (iii) 0-56.

The small size of the soldier of this species will at once separate it from any other
described Australian Hamitermes ; its nearest ally is Hamitermes latidens, Mjob.,
from which it may be distinguished, z¢fer alia; by its smaller size and the form and
number of joints in the antennae.

NORTH QUEENSLAND : Townsville.

Biology.

Winged adults, soldiers and workers were found in heavily manured garden soil
on 10th December and 18th December 1919. About 50 points of rain fell on the night
of 17th December, and at 8.30 a.m. on the following morning winged termites were
noticed flying up from the very sandy soil. After some searching a small circular
opening was found on a bare sandy space from which the winged forms were
emerging. One soldier and several workers were congregating about the opening,
apparently guarding the entrance to the nest while the winged forms made their
exit. Between 8.30 a.m. and 9 a.m. on 16th and 17th November 1920, winged forms
were again noticed rising from the grass-covered soil a few yards distant from where
the first specimens were taken in the previous year. In most cases they fluttered
feebly from a blade of grass or other object, flew a few yards and settled again in the
grass, where most of them were caught almost at once by small ants (Pheidole
megacephala). The first swarm (16th November) was preceded a few hours earlier
by a heavy shower of rain and the second swarm, firstly, by a very copious watering
of the adjacent soil from the pipe service and, secondly, by a heavy shower of rain
about 5 a.m.

Hamitermes perplexus, sp. n.

Imago.—Head and prothorax dark brown (bay); postclypeus lighter than
head, anteclypeus whitish with yellow blotches ; antennae and tergites of abdomen
mummy brown ; on the first nine tergites a small clear mark on each side, distinctly
comma-shaped on segments 3-6 inclusive; sternites Dresden brown, first four
paler than others and only very slightly darker at sides, the rest uniformly dark,
plates 1-6, inclusive, with small clear mark at each end, 5-8 uniformly dark ; sternum
and pleurae not darker than sternites of abdomen ; wings brown.

Fig. 16. Hamitermes perplexus, sp. 0.,
head of imago.

Head (fig. 16) rounded behind, flat on summit, widest across the eyes, moderately
hairy, hairs of variable length. Eyes large, projecting beyond sides of head. Ocelli
oblique, oval, widely separated from the eyes. Fontanelle large, oval, with linear

(5296) 2F2

382 GERALD F. HILL. |

forward prolongation, two oblique impressions in front. Anteclypeus produced in
front ; postclypeus large, convex, rounded behind, truncate in front, with distinct
median suture. Antennae 15-jointed; Ist twice as long as 2nd; 3rd very short,
smallest of all; 4th and 5th nearly equal to each other, larger than 3rd; 6th larger ©
than 5th; 7th-12th nearly equal; 13th and 14th very little longer and more cylin- |
drical than 12th; 15th longest, tapered from before the middle to the pointed tip.
Very rarely the 5th and 6th joints on one side only are fused.

Prothorax slightly bent up in front, antero-lateral margin slightly rounded,
sides narrowed to the emarginate posterior border, divided medially by a suture,
which is very distinct in the anterior half ; two clear impressions on each side behind —
the anterior margin; the whole surface moderately hairy. A distinct pale mark —
on each side of the median line of the metanotum.

Wings (fig. 17) with border ciliate excepting on proximal fifth of hind margin ; |
venation very variable, often differing in both fore- and hind-wings of same insect ;
radius darker than costa, sometimes with short branches near apex of wing. In_
the fore-wing the median branches from the radius within the wing-stump, in the hind- _
wing the division is well beyond the suture ; the median runs nearer to the cubitus _
than to the radius and gives off from three to eight branches to the anterior margin, |

Vig. 17. Hamitermes perplexus, sp. n., wings
of imago.

apex, or to the hind margin, the first branch generally arising about the distal third |
or fourth of the wing, but sometimes about the middle of proximal third. In some >
a short stout vein branches from the median at the proximal fifth and joins the radius |
about its proximal fourth; cubitus very irregular and seldom alike in the fore- or
hind-wings of the same individual. In the fore-wing there are generally from six |
to eight simple branches, all of which often reach the posterior border before the |
middle ; in the hind-wing there are from seven to eleven branches, some of which are
forked. Associated with individuals having a wing venation as described above |
there are generally many with the venation described by Mjéberg (1920, p. 84) in_
H. obtusidens, Mjob. |
|

Legs ochraceous tawny, short; tibial spurs, 3:2: 2.
fo) p)

Abdomen with ten distinct dark-coloured tergites; ventral surface much lighter. |
Styli wanting in both sexes. |

Measurements: Length with wings, 12-13; length without wings, 7-8; head,
with mandibles, 1-27 long ; thorax and abdomen, 6-25 long ; head, including eyes,
1-17 wide; mandibles, right, 0-72 long, 0-51 wide, left, 0-73 long, 0:41 wide; |
antennae, 1-7-1-8; prothorax, 0:61-0:65 long, 1-1-17 wide; tibia (1) 0:85-0:89, |
(ii) 0-84, (iii) 1-17; abdomen, 1-9 wide.

Neoteinic Queen.—Head and thorax ochraceous tawny to light orange-yellow ; |

tergites of abdomen paler, rest of insect creamy. |

ae ee rl

ON SOME AUSTRALIAN TERMITES. 383

Head about as long as wide, widest at the eyes, hairy. Eyes small, not pro-
jecting, pigmented in centre. Ocelli roundish, well separated from the eyes. Fon-
tanelle a large globular protuberance. Anteclypeus half as long as postclypeus,
nearly truncate in front, sometimes concealed; postclypeus convex, depressed in
the middle line, hairy. Labrum swelling out on the sides, bluntly rounded in front,
not covering apical teeth. Antennae 15-, rarely 16- or 17-jointed, very variable.

Prothorax as wide as head, shaped as in imago, clothed with red hairs. Wing-
pads short, hind pair not reaching beyond middle of first tergite.

Measurements: Total length, 9-5-10; head, with mandibles, 1-32 long, 1-12
wide ; abdomen, 2-8-3 wide.

Soldier —Head ochraceous ; mandibles pale ferruginous, a little paler at the
base ; antennae ochraceous with pale segmentations ; clypeus and labrum stramineous
with ochraceous blotches, remainder of insect stramineous.

Fig. 18. Hamitermes perplexus, Fig. 19. Hamitermes germanus,
sp. n., head and prothorax of Hill, head of soldier.
soldier.

Head (fig. 18) longer than wide, slightly curved on the sides, a bright ferruginous
spot on either end of the clypeus. Labrum large, rounded on the sides, rounded
in front, not reaching the mandibular teeth. Clypeus wider than long, convex,
emarginate in front, divided in the median line by a deep cleft which extends into
thehead. Mandibles long and moderately slender, falciform, with a sharp backwardly
directed tooth about the middle similar to that of H. germanus, Hill (fig. 19).
Antennae 15-jointed; Ist joint twice as long as 2nd; 3rd half as long as 2nd,
smallest ; 4th and 5th nearly equal in length; 6th and 7th longer than Sth.

Prothorax (fig. 18) saddle-shaped, rounded and bent up in front, antero-lateral
margin rounded, posterior margin rounded, slightly emarginate in the middle.
Legs rather short and stout; tibial spurs, 3:2: 2.

Abdomen with scattered, rather short, and a few long yellow hairs. Cerci with
base short and moderately broad, apex elongate, sides nearly straight to near the
pointed tip.

384 GERALD F. HILL.

Measurements: Total length, about 5; head and mandibles, 1-83-1-97 long ;

thorax and abdomen, 3-1-3-7 long ; head, 1-08 wide, 0-84 deep ; mandibles, 0-98 —

long; antennae, 1-69; thorax, 0-36 long, 0-7 wide; tibia (i) 0-84, (ii) 0-75, (ii) 1-08.
Worker.

Head creamy ; thorax and legs paler; antennae whitish, tinged with |

yellow-ochre, distal third darkest ; abdomen white (greyish black when alimentary —

tract contains ingested matter).

Head about as long as wide, clothed with reddish hairs, some moderately long. —

Labrum large, covering mandibles. Anteclypeus narrow, bluntly pointed in front ;

postclypeus convex, with reticulate pattern. Mandibles with dentition as in imago. ~

Antennae 15-jointed ; Ist joint large, nearly twice as long as and much wider than

2nd; 3rd very small, smallest ; 4th and 5th about equal to each other; 6th larger

than 5th, but much smaller than 7th and 8th.

Prothorax with anterior portion narrowed and bent up, emarginate in the middle,
posterior margin almost semicircular, the whole clothed with reddish hairs of unequal
length.

Abdomen elongate-oval, moderately hairy. Cerci small, basal part without hairs,
apical part tapered and drawn to a fine point at tip.

Measurements: Total length, about 5-15; head, with jaws, 1-15 long ; thorax
and abdomen, 4; head, 1-5 wide; mandibles, right, 0-61 long, 0-51 wide, left, 0-66
long, 0:42 wide; antennae, 1-46; prothorax, 0-37 long, 0-8 wide; tibia (i) 0-76-
0-78, (11) 0:67-0-7, (iii) 0-99-1.

NORTH QUEENSLAND: Townsville.

This species appears to be closely related to H. laurensis, Mjéberg, from which
it differs in the shape of the head and antennae of the soldier and in the antennae
of the neoteinic queen. From H. mevidionalis, Frogg. (fig. 20), it differs in the

Fig. 20. Hamitermes meridionalis,
5 .
Frogg., head of soldier.

soldiers in the shape of the jaws, colour of the head, form of the antennae, shape
of the clypeus and distance of the antennae from the head margin. The winged
forms of H. laurensis and H. meridionalis are not known.

ON SOME AUSTRALIAN TERMITES. 385

Biology.

This is one of the commonest, if not the commonest, species of termite found
in the vicinity of Townsville ; how much further it ranges is not known. Like all
the species of this genus known to me it is not a wood-eater, but lives on vegetable
debris, such as the leaves, stems and seeds of herbaceous plants, the two former being
cut into fragments and the latter stored intact in the termitaria. ood is gathered
throughout the year, as required, and is stored in the outer galleries. In times of
drought or after grass fires there is a noticeable absence of food in these galleries,
and at no time is the accumulation of food comparable with that which is found in
the nests of the grass-eating species of Eutermes. Incipient colonies, and colonies
which for some reason or other have no termitaria, do not appear to make any
provision for the storage of food in their underground galleries.

In these notes the word termitaria is used to indicate earthy or woody nests
constructed by termites upon the surface of the soil, upon stumps, logs, stones,
or in the branches of trees; the latter—arboreal nests or termitaria—appear to be
constructed only by certain species of Eutermes. The species under notice normally
constructs earthy nests, but it appears that these are not commenced until the colony
has matured a considerable number of individuals of the worker caste in under-
ground galleries. In some cases winged adults also are reared in these galleries.
In discussing the habits of H. eucalypti, sp. n., reference is made to the construction
of tube-like vertical ducts, by means of which the winged adults leave their under-
ground galleries at the time of the annual colonising flight or “swarming.” — Similar
ducts are also, though rarely, constructed by H. perplexus for the same purpose,
when winged adults are reared in underground galleries. These tower-like ducts are
commenced at the end of the dry season after the first showers and before the regular
rain sets in; for example, in the summer of 1919 the first heavy shower (0-5) fell
on 17th December, and construction commenced on the following day and continued
until 22nd December, by which date about 80 ducts were scattered over an area
of about 20 ft. square.

It is acommon occurrence to find small colonies of soldiers and workers in rambling
galleries in the soil, or at the base of abandoned, or partly abandoned, termitaria
of Eutermes and Drepanotermes, and in one instance a de-alated imago of Eutermes sp.
was found with such a community.

Typical termitaria, however, are very commonly found on hillsides, near the foot
of hills (Pl. x, fig. 1), and on the open or scrub-covered plains (PI. x, fig. 2) to the
westward of the town, and occasionally in town gardens and streets. Quite
frequently the greater part of the mound rests upon a large rock projecting a few
inches to a foot or more above the surface. In such cases, however, one portion of
the mound is invariably in direct contact with the soil, and is pierced by the main
galleries communicating with the earth below. It may be stated here that such
communication is absolutely essential to the existence of all termites, excepting
members of the genus Cryptotermes, which in many, if not most, cases normally live
entirely cut off from access to the soil. By far the greatest number of nests are to
be found on the open grazing country and adjacent scrubby areas. In the former
localities the nests attain their maximum size and number, and associated with them
are almost as many nests of Eutermes sp. and Drepanotermes silvesivit, sp.n. Near
the margin of the lowest lying part of this area the nests of these three species are
sO numerous as to present a most remarkable feature of the landscape.

In size and shape the nests vary a good deal ; those on higher and well drained
positions being usually smaller and more pointed than those on the plains (Opn lees
figs. 1 and 2). In the ground-plan, nests on the higher situations are circular or
oval, in the latter case the long axis being directed north andsouth. A nest measuring,
say, about 12 by 14 in. at the base would be about 15-18 in. high, with sides sloping

385 GERALD F. HILL.

to the pointed apex. On the plains the nests are often very much larger and

frequently they have the long axis directed north and south or north-west by south-
east, the sides sloping to the bluntly wedge-shaped top, which is rounded off at each
end and never surmounted by numerous small points as in H. meridionalis, Frogg.
(Plate xi). Sometimes the western side is more or less convex and the eastern side
vertical or bent over, asin the latterspecies. The maximum size of sucha termitarium
is about 6 ft. long by 2 ft. wide at the ground and 4 ft. high. Conical nests are not
uncommon.

Only in recently constructed nests, built by large colonies whose former nests
have been dismantled, or smaller original nests of strong colonies, are the galleries
and passages within the termitarium numerous and extensive. In the great majority
of cases the structure is intensely hard and composed almost entirely of earthy
particles cemented together. The occupied portions are practically confined to the
top and sides, the internal galleries being gradually filled with rejectamenta until
they assume the toughness and density of the earthy portions. Additions are made
throughout the year, as in H. meridionalis, and not only after the rainy season has
set in, as stated by Jack (1897). These additions are small and local and usually
take the form of thin layers added to the sides (Pl. xu, fig. 2), the insects working
from holes cut in the adjacent walls. The almost solid interior is pierced by a few
larger passages which pass down into the soil and beneath the walls. Food is stored
in the outer galleries and is generally mixed with the bodies of their dead. Near the
top of the nest the dead occupy more space than does the vegetable food. They

consist chiefly of nymphs of the winged forms in the stage when the wing rudiments |

first appear ; but workers and soldiers are to be found also. In nearly all cases the

legs have been amputated. This habit of storing away the dead was first recorded |
by Mjéberg (1920) in H. laurensis, Mj6b., H. meridionalis, Frogg., and Eutermes |

tyriet, Mjéb., in North Queensland, but has long been known to occur in H.

meridionalts in the Northern Territory, and in an allied species in Central Australia. |
Another type of termitarium is that in which there is a more or less solid |

foundation a foot or more in height, upon which rest several larger or smaller cone-
shaped points (PI. xu, fig. 1). In many instances these nests are obviously constructed
on the sites of old nests ; in others there is nothing to indicate that such is the case.

An examination of the smaller nests, and especially those constructed on land free |

from rock, shows clearly that the base rests on the natural surface, which is
penetrated only by a few passages. Such nests are easily pushed over intact, and if
not removed or broken into fragments, form the base of new termitaria, which the
insects soon construct in the characteristic cone-like form. Sometimes several of
these cones are built up vertically upon the upper surface of the now recumbent
old structure.

The termitarium invariably contains two sterile castes, namely, soldiers and

workers. In all species of the genus Hamitermes (sensu restricto) the former caste |
is represented by very few mature individuals, probably never more than 5 per cent. |
of the total number of workers, and in this species probably less than } per cent. |
When the nest is broken into there is a general retreat to the remaining galleries, |

neither caste making any attempt to defend themselves, their fellows, or their home.

In all strong colonies there are present also great numbers of young forms in various |

stages of development. Reproductive forms, or forms which mature into them, are
nearly always present. In the earlier stages preceding sexual maturity the latter

are creamy white or white, soft-bodied insects, longer and more slender than the |
workers and possessing short wing-rudiments or wing-buds.* These nymphs are |
often present in great numbers and appear to be destroyed and stored for food when |

* The term ‘‘nymph’”’ is used in this paper to denote the young of reproductive forms in

which the wing rudiments are evident. The word “larva” is used to denote all apparently

undifferentiated young.

ON SOME AUSTRALIAN TERMITES. 387

produced in excess of requirements. The survivors may develop into sexually mature
insects of four kinds, namely, neoteinic, or supplementary, kings and queens or true
kings and queens. In the ‘former (neoteinics) the wing-buds undergo very little
further development, and the eyes and chitinous parts become only partly pigmented.
After fertilisation the abdomen of the female becomes greatly enlarged, and she is
capable of laying a great number of eggs, which, in this species at any rate, produce
individuals of all castes. Those nymphs that are destined to develop into true
kings and queens continue to develop the wing-buds until, after a final moult, they
emerge as winged imagines, with pigmented eyes and chitinous parts.

In all of the many scores of termitaria examined, the reprodtfctive forms found
in each were either neoteinics only, or one true king, and from a few to over 100
neoteinic queens. True queens have not been found, nor have neoteinic kings been
observed in any nest in which a true king was present; there are no queen cells or
nurseries in these termitaria, the ordinary flattened horizontal cells being utilised
as required. The ovigerous neoteinic queens are to be found scattered through
all parts of the nest, but generally near the walls. Each queen appears to preside
over a restricted area, in which the eggs and young larvae are to be found, the latter
in one or more small clusters about the size of a large pea. As the larvae develop
they spread out in all directions, mixing with the soldiers and workers. The king
is rarely associated with one of these queens ; more often he is to be found in one of
the larger cells with from 10 to 40 younger queens of various sizes, some only
recognisable as such, others apparently as fully developed as the egg-laying individuals.
Neoteinic kings, when present, occupy these cells and have not been found i in cells
occupied by isolated queens. It appears that, as a rule, queens are fertilised and
attain nearly their maximum development before they migrate to other parts of the
nest to oviposit, and that they are not often re- fertilised from time to time, as is
believed to be the case with true queens of other species. The fact that old neoteinic
queens, 7.¢., queens with shrunken abdomens, have not been found in this species
and that males are very rarely found with isolated queens, suggests that normally
a queen does not mate after she begins egg-laying and that she is destroyed and re-
placed by a more fecund one as soon as she has passed her prime.

Prior to the final moult, which takes place about November, pigmentation in the
nymphs of the first form is ‘confined to the eyes. The wing-buds are short and thick
outgrowths from the posterior margin of the meso- and metathorax, measuring
about 3 mm. in length. After the moult the wings appear as soft, white, crinkled
membranes, which rapidly assume their full length, Ga remain unpigmented, like
the rest of the body, for some days. The dur ation of the period intervening between
the moult and acquisition of the full degree of pigmentation of the chitinous parts
and functioning wings is not known, but it appears to be about 10-14 days. Moulting
does not take place “simultaneously in all the individuals which are destined to take
part in the colonising flight, many being still in the final nymphal stage while others
are capable of flight. Indeed, some individuals do not develop functioning wings
until after the flight. W hether the true king, which is so often found with the
neoteinic queens in the termitarium, is developed from one of these, or from one
of the earlier matured imagines, or whether he is the original male parent of the colony,
is not known. The latter, however, is most improbable, since he rarely, if ever,
possesses blunted claws and pigmented apices to the tarsi, which are the indications
of age in mature queens. Further, many of the termitaria are evidently very much
older than the longest period suggested as the probable life of this caste.

The life of the winged imago within the termitarium is short, and is certainly
regulated to a consider able degree by weather conditions, that is, swarming does not
occur before the first heavy rain of the season has fallen and the ground is thoroughly
moistened.

388 GERALD F. HILL.

It has been stated above that true kings, 7.e., de-alated, sexually mature imagines,
are often found in the nests as consorts of neoteinic queens, but true queens are
unknown in this species. The questions that naturally arise are: What becomes of
the countless thousands of female imagines (potential queens) which issue yearly
from most, but not all, termitaria 2? Do none of them become the founders of new
colonies, as is the case with many species? Is this species perpetuated solely by
neoteinic queens and true kings or neoteinic kings? It is beyond doubt that the
vast majority of individuals of a colonising flight, 7.e., potential true kings and
queens, succumb to the attacks of predacious ants, lizards and birds within a few
minutes of the commencement of their free life, but it is hardly conceivable that all are
irretrievably lost. Observations made during a period of two years, during which
many scores of nests have been examined at fairly frequent intervals, convince me
that the normal manner of reproduction is by the forms commonly found in the
nests, 7.e., true and neoteinic kings and neoteinic queens. In seeking an explanation
for the apparent non-existence of true queens, one suggests itself as being the most
probable, namely, that the life of the winged forms, and probably of the workers
and soldiers also, is short, perhaps about two years in the case of the former, that
a small proportion escape destruction at the time of the colonising flight, mate and
become the parents of the colonies which for some time live in galleries in the soil.
Later, these colonies increase to such size that the construction of a termitarium
becomes possible, or perhaps necessary; in the meantime, however, the founders
have lived their lives and their place in the community has been taken by neoteinics
derived from nymphs. This theory does not account for the presence of young true
kings in old termitaria, except in the manner suggested in a previous page, nor does
it explain why, if male imagines are sometimes retained in the community to be the
consorts of neoteinic queens, female imagines also are not retained to obviate the
necessity for bringing neoteinic queens into use. Explanations of these and other
phenomena in the economy of termites present difficulties which can be overcome only
by prolonged and careful field observations.

The invasion of these termitaria by the common ant, Jvidomyrmex sanguineus,
Forel, has been referred to in discussing the biology of Drepanotermes silvestri, sp. 0.

Hamitermes perplexus, var. victoriensis, nov.

King.—Colour as in H. perplexus, sp. n., excepting that the antennae and tergites
of the abdomen are lighter (Dresden brown) ; sternites Dresden brown, the first
five distinctly darker at the sides than elsewhere, 6th-8th uniformly dark, no pale
marks visible ; the first seven tergites only have pale marks at each side, more or
less indistinct excepting on 3-6 inclusive; sternum and pleurae argus brown.

Head as in H. perplexus, excepting as follows: Ocelli smaller, fontanelle smaller,
linear extension rather longer. Thorax as in H. pferplexus, except that the two
smaller lateral impressions on prothorax and the clear marks on metathorax are
wanting. Legs as in H. perplexus. Abdomen as in H. perplexus, excepting as noted
above. Measurements as in H. perplexus.

Neoteinic Queen.—Head, thorax and wing-stumps light orange-yellow, sternites
and tergites of abdomen of the same colour ; remainder of abdomen and legs creamy ;
the whole insect clothed with fine, pale hairs.

(2) Head longer than wide. Eyes small (0-18), pigmented in centre. Ocelli
small, situated as in king. Antennae 14-jointed, segmented as in king. Fontanelle
not a depression, but a broadly lanceolate scar, with a short straight median line.
Thorax as in king, wing-pads of metanotum extending to the middle of the third
tergite. Legs with tibial spurs worn down to short, blunt stumps. Apices of the
first three tarsals heavily chitinised. Styli absent. (This appears to be a normal
queen of the second form.)

Measurements: Total length, 12; head, with mandibles, 1-5 long, 1-22 wide ;
prothorax, 0:65 long, 1-12 wide; abdomen, 3-25 wide.

ON SOME AUSTRALIAN TERMITES. 389

(4) Gross appearance similar to that of (2). Labrum very short, exposing the
two apical teeth of both mandibles. Anteclypeus invisible. Eyes and ocelli as in (a).
Antennae 15-jointed. Fontanelle a large cone-shaped projection. Wing-pads of
metanotum extending to the base of the second abdominal tergite. Styli absent.
(This is evidently an abnormal form.)

Measurements: Total length, 7-25; abdomen, 2 wide.

Soldier very similar to H. perplexus. Labrum slightly more rounded; clypeus
rather narrower and more deeply emarginate ; antennae inserted nearer the outer
margin of head, first two joints not so long and slender ; mandibular teeth rather
more hook-like ; gula as seen in dotted line (fig. 21).

Fig. 21. Hamitermes perplexus
var. victoriensis, nov., head of
soldier.

Measurements : as in H. perplexus.
Worker.—As in H. perplexus.
VICTORIA: Preston (F. E. Wilson).

The differences between the Victorian and Townsville specimens are very slight
indeed, but appear to me to be sufficient to justify one in separating the former as
a variety of the latter.

Biology.

This variety is described from two small colonies found under stones. The first
colony was taken on 14th November and comprised a few larvae, soldiers, workers,
nymphs of the second form and the neoteinics described above. The eyes of the
nymphs are small and very little pigmented, the ocelli are rudimentary, the antennae
15-jointed as in the imago, and styli are present in both sexes. Worker-like indi-
viduals of whitish colour and slender form I consider to be immature reproductive
forms in the stage prior to the development of the wing rudiments.

The second colony was taken in September and comprised larvae, soldiers, workers,
nymphs of the second form and a true king. Neither a true nor a neoteinic queen
was found. It appears to be a rare form and the first of the genus to be recorded
from Victoria. There are no termitaria of any kind in the district.

390 GERALD F. HILL.

Hamitermes neogermanus, sp. n.

Soldier.—Head and base of mandibles orange, mandibles ferruginous at the tip ;
antennae mars yellow, darker than head ; remainder of insect pale stramineous.

Head and mandibles (fig. 22) together twice as long as wide, rounded behind and
on the sides. Mandibles very long, falciform, with a sharp hook-like tooth before
the middle. Labrum almost as long as wide, rounded in front. Clypeus large, wide,
with median shallow depression not reaching the posterior margin. Antennae
15-jointed, long and slender ; 1st joint long and narrow, twice as long as 2nd; 2nd
cylindrical; 38rd short and narrow, smallest of all; 4th and 5th alike, about twice
as long as 3rd; 6th longer, nearly as long as 7th; 7th-12th alike; 13th and 14th
a little longer ; 15th rather longer than 14th, bluntly pointed.

Prothorax shaped as in H. ferplexus, sp. n., with scattered long red hairs. Legs
moderately slender, similar to those of H. perplexus, fore-tibiae with a row of long and
stout hairs on the lower side of the apical half.

Fig. 22. Hamitermes neogermanus,
sp. n., head of soldier.

Abdomen wide in the middle, tapered to the pointed apex. Cerci short.

Measurements: Total length, 5-5; head and mandibles, 2-16 long; head,
1-17 wide; mandibles, 1 long; antennae, 2:03; thorax and abdomen, 3:5 long ;
prothorax, 0-42 long, 0-7 wide ; tibia (i) 0-92, (ii) 0-84, (iii) 1-17.

Worker.—Head and thorax pale stramineous ; antennae much darker towards
the apex ; remainder of insect whitish, clothed with red hairs.

Head rounded behind, rather straighter on sides than usual. Labrum large,
narrower at the base than across the middle. Anteclypeus large, produced in front.
Postclypeus large, convex, truncate in front, rounded behind. Antennae 15-jointed ;
Ist joint twice as long as 2nd; 3rd very short and narrow, smallest ; 4th and 5th
about equal to each other; 6th larger ; 7th—15th increasing in length gradually.

Prothorax with anterior portion rounded and bent up, antero-lateral angles

rounded, posterior margin rounded, with slight emargination, clothed with long
and short red hairs. Legs rather short and slender.

Abdomen elongate-oval, clothed with red hairs. Cerci with basal portion short
and broad, remaining two-thirds slender.

ON SOME AUSTRALIAN TERMITES. 391

Measurements: Total length, 5-25; head, with mandibles, 1-5 long; head
1-12 wide ; antennae, 1:41 ; thorax and abdomen, 3-76 long ; prothorax, 0-37 long,
0-84 wide ; tibia (i) 0-86, (11) 0-7, (iti) 1 ; abdomen, 1-8 wide.

This species is most closely related to H. germanus, Hill, from the Northern
Territory (fig. 19); it differs, however, in its much larger size, larger and more
rounded labrum, and larger and differently shaped clypeus. In both species the
antennae are similarly segmented and the mandibles are alike except in size.
The two species are easily separated.

Described from one soldier, and numerous nymphs and workers in alcohol, and
nine soldiers and nine workers on cards.

Type series in South Australian Museum ; co-types in author’s collection.

SouTH AusTRALIA: Mt. Lofty Ranges (N. B. Tindale); Gawler (A. M. Lea);
Angaston (A. M. Lea).

Hamitermes eucalypti, sp. n.

Imago.—Palpi and legs buckthorn-brown; head, thorax, wings, abdominal
tergites mummy-brown, head darkest, antennae Dresden brown, darker towards
the tip.

Head (fig. 23) rounded, slightly longer than wide, widest across the eyes, densely
clothed with moderately short hairs. Maxillary palpi with first and second joints
very short, together equal to the third; third and fourth equal. Eyes large,
rounded, projecting well beyond sides of head. Ocelli large oval, separated from the
inner margin of the eyes bya distance equal to their length. Fontanelle large,
broadly lanceolate, twice as long as wide.

Fig. 23. Hamitermes
eucalyptt, sp. n., head
of imago.

Antennae 15- or 16-jointed, dark with clear articulations; 1st joint twice as
long as 2nd and much wider; 2nd cylindrical and about as long as 3rd and 4th
together ; 3rd very short, narrower than 2nd and 4th, indistinctly separated from
the latter; 5th and 6th equal in length and shorter than 7th and 8th; 9th to 13th
longer than 7th and 8th and nearly equal to each other ; 14th slightly longer than
13th ; 15th and 16th a little longer than 14th; 15th cylindrical.

Prothorax slightly bent up in front, rounded on the sides to the truncate posterior
margin, a clear oblique oval area on each side of the median suture in the anterior
third, the whole surface moderately densely clothed with medium-sized hairs. Wing-
stumps small, triangular, clothed with numerous short and long hairs, suture straight ;

392 GERALD F. HILL.

margin of wing (fig. 24), excepting proximal fourth of posterior border, ciliate, with
scattered hairs on membrane, especially on apical half and along veins; costa, radius, _
base of median and first seven or eight branches of cubitus very distinct, branches
of cubitus sometimes anastomosing to form cells. Legs with femora moderately
stout; tibial spurs, 3:23 2.

Big. 24. Hamitermes eucalypti, sp. n.,
wings of imago.

Abdomen long and nearly cylindrical, with 10 dark tergites, each of the first
eight very distinct and with a small clear spot towards each lateral margin, moderately
hairy ; eight distinctly visible sternites, pale except at the lateral margins, where
there is on each side a dark spot, sixth longest and darkest. In the male there are
seven distinctly visible sternites, with lateral spots on the first three, the remainder
nearly uniform in colour; lower surface darker than in female. Cerci short, basal
portion as long as apical.

Measurements: Length with wings, 11-12; length without wings, 6-5-8;
head, with mandibles, 1-2-1-3 long; head, at and including eyes, 1:1 wide;
mandibles, right, 0:-6-0-62 long, 0-46 wide, left, 0:65-0:67 long, 0:35-0:37 wide ;
antennae, 1-65; prothorax, 0-47 long, 0-98 wide ; fore- and hind-wings, 9-25 long ;
fore-wing, 2:58 wide; hind-wing, 2:68 wide; tibia (i) 0-7-0-75, (ii) 0-8,
(iii) 1-1-1-13.

Fig. 25. Hamitermes eucalypti, sp. n., head of
soldier (a) from above; (bd) in profile.
Soldier.—Head ochraceous, mandibles ferruginous, labrum whitish; thorax
lighter than head; abdomen and legs very light yellow.
Head with mandibles (fig. 25) longer than wide, rounded behind and on the sides,
with scattered red:hairs. Labrum convex, widest at the base, sloping to the bluntly

ON SOME AUSTRALIAN TERMITES. 393

pointed apex. Clypeus convex, much wider than long, arcuate in front, divided by
a deep median suture which extends into the front of the head, where there is a
flask-shaped depression. Mandibles long and moderately slender, falciform, with
a sharp hook-like tooth near the middle. Antennae 15-jointed, arising from a
prominence within a short and wide fossa behind the base of the mandibles ; Ist
joint very large, 2nd less than half as long as Ist and much narrower ; 3rd very short
and narrow; 4th and 5th equal in size, longer and wider than 3rd; 6th much longer
and wider than Sth.

Thorax with anterior half bent up, lateral margins angular, posterior margin
rounded, clothed with scattered reddish hairs as on head. Legs with femora
moderately stout, fore-tibiae stout, mid-tibiae with two short stout setae towards
the apex in addition to the two larger apical spurs ; tibial spurs, 3:2: 2.

Abdomen clothed rather densely with reddish hairs.

Measurements: Total length, about 4; head, including mandibles, 1-78-1-8
long; thorax and abdomen, 2-8 long; head, 1:03-1:12 wide; antennae, 1-41 ;
prothorax, 0°32) long) 0-% wide; tibia (i) 0-8) (11) 0+7, (ui) 1; abdomen, 1-17
wide.
| Worker.—Head and thorax creamy, rest of insect almost hyaline, the whole
_ surface moderately hairy.

Antennae 15-jointed ; Ist joint twice as long as and one-fifth wider than 2nd ; -
_ 8rd very short and narrow, shorter than broad ; 4th and 5th equal in length, longer
and wider than 3rd; 6th as long as 2nd; 7th—12th similar to each other, longer than
6th ; 13th and 14th equal in length, the latter more cylindrical ; 15th as long as Ist,
widest at the proximal third, tapered to the pointed apex.

Prothorax with anterior third rounded and bent up, emarginate ; lateral margins
elongate, bluntly pointed; posterior margin truncate, without emargination ;
surface clothed with stout reddish hairs. Legs similar in shape and armature to those
of soldier, except that the two short setae on the upper surface of the second tibiae
of the soldier are replaced by much longer and more slender ones ; tibial spurs,
O22.

Measurements: Total length, about 5-2; head, with mandibles, 1-41 long ;
thorax and abdomen, 3-76 long; head, 1-04 wide; mandibles, right, 0-51 long,
0-42 wide, left, (a) 0-51 long, 0-3 wide, (b) 0-61 long, 0-37 wide; antennae, 1-4-1-7;
prothorax, 0-37 long, 0-75 wide ; tibia (i) 0-7, (ii) 0-65, (11) 0-90-0-93.

This species appears to be nearest H. herbertensis, Mjéb., from which it is easily
distinguished by the characters given for the imago and soldier.

N. QUEENSLAND: Magnetic Island, Townsville.

Biology.

This is a common species in the Townsville district and on Magnetic Island,
where it is generally found under earthy covered-ways on the trunks of living
eucalyptus trees growing on hill-sides and open forest lands. Very rarely these
covered-ways are constructed on the trunks of dead trees from which the bark has
fallen ; but in most cases living trees with friable bark are favoured. At first, tube-
like covered-ways are constructed, which extend up the trunk to a height of 6-10 ft.,
the loose, weathered surface being removed as the tube progresses upwards. Later
on these tubes are extended laterally until a considerable surface is encased. On
removal of this fragile casing the outer weathered surface of the bark beneath will
be found to have been removed—apparently for food. On one occasion a very
small and inconspicuous mound of earth, unlike a true termitarium, lay at the base
of the tree, and from it the casing extended up the trunk for a distance of 33 ft.,
where it had been cut off. The mound was traversed by a few galleries, which con-
tained a few soldiers, workers, and nymphs of the second form (18.viii.1919).

394 GERALD) F. HILL.

The trunk, which was hollow, was sealed up on top with earth. When a portion
of the mood was cut away, the interior was found to be nearly filled with earthy
material, pierced by a few galleries, which contained the castes found in the mound.
Four months later (15.i.1920) Coptotermes (?)lacteus, Frogg., was found in possession

of the trunk a had commenced to build a typical termitarium on the site formerly

occupied by the small mound referred to above. H. eucalypti was found under the
casing on ae bark and in tunnels in the adjacent soil; in the latter were
also three winged adults. It appeared that the invading Coptotermes were the
survivors from a large termitarium 6 ft. distant, which had been destroyed on
{8th August 1919.

Fig. 26. Vertical tubes constructed by Hamiterimes eucalypti,
sp. n., prior to swarming.

On 25th November 1920, at Magnetic Island, near Townsville, the trunk of a large
bloodwood tree (Eucalyptus) was found to have its sunny side almost encased in
an earthy crust, under ie h were found many workers and a few soldiers of this
species. Nearer the ground a few adult winged forms were found. Arising from the
sandy soil near the trunk and within a space of a few feet were about a dozen vertical
tubes (fig. 26), ranging from 5—23 cm. in height by about 1-3 cm. in diameter, which
communicated each with an irregular ch amber about 25 cm. long by 14 cm. wide,

VOL. XII. Part I.—pp. 1-106.
JUNE, 1921.

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ON SOME AUSTRALIAN TERMITES. . 395

and these again with a system of small tunnels extending downwards to a depth of
about 25 cm. Workers and soldiers only were found in the tubes, but numerous
imagines and a few nymphs with short wing-pads were collected in the chambers and
galleries. The latter appeared to be nymphs of the second form, 7.e., nymphs which
would mature into queens and (?) kings of the second form (supplementary royalties).
On 4th December a few winged adults were captured at a lamp in this locality.

From my observations I conclude that termites of this species do not construct
termitaria, but rear their young and reach maturity in underground galleries. The
vertical tubes and adjoining chambers are constructed just before the winged forms
reach maturity and for the purpose of enabling these forms to assemble and disperse
unmolested by predatory ants and other ground-frequenting enemies. The advantage
of these tubes will be apparent to anyone who has witnessed the decimation of the
winged forms of certain other species, which make their exit at the time of swarming
by an opening made in the surface of the soil. Elsewhere in this paper reference
is made to similar tubes constructed by Hamitermes perplexus, sp. n., which
normally builds termitaria, but which frequently lives in underground galleries, as
does H. eucalyptr.

Leucotermes clarki, sp. n.

Imago.—Upper surface buckthorn-brown, seven hindmost abdominal tergites
a little lighter ; mouth-parts and lower surface testaceous ; abdominal sternites not
distinctly darker ; wings brown, apical half near costa and radius darkly shaded.
The whole insect densely clothed with short reddish hairs.

Head (fig. 27) large, wider than long, widest at eyes. Fontanelle elongate,
widest behind, two oblique clear impressions on either side in front midway between
anterior end of fontanelle and posterior margin of clypeus; an indistinct suture
extending posteriorly from fontanelle. Eyes very large, circular (0-28 dia.), projecting
well beyond sides of head. Ocelli large, nearly circular, half as wide as eyes, from

Fig. 27. Leucotermes clarki,
sp. n., head of imago.

which they are separated by a rather narrow space. Anteclypeus very small, one-
fourth as long as postclypeus, whitish, produced in the middle. Postclypeus large,
paler than rest of head, darker than antennae, convex above, slightly curved in front,
- more so behind, twice as wide as long, divided medially by a narrow dark suture.
Antennae 15- or 16-jointed; in 15-jointed antennae, Ist stout, twice as long as and
much wider than 2nd, 3rd smallest, 4th and 5th equal, 6th-12th increasing
slightly in length, 13th longer, 14th about as long as 13th but more cylindrical,
15th elongate-oval; in 16-jointed antennae the 8rd joint of the other form is
divided into two, the first joint is generally more slender; the 3rd joint is very
small and fused with the 4th, which is only a little larger.
(5296) 2G
»

396 GERALD F. HILL.

Thorax moderately flattened above, rounded, slightly bent up and emarginate
in front, rounded on the sides, narrowed to the shghtly emarginated posterior border,
divided medially by a suture, two clear transverse impressions behind the upturned
anterior margin. Legs slender; tibial spurs, 3:2: 2.

Wing-stumps triangular, suture straight, margin ciliate except at base of
posterior border, scattered hairs over entire surface of membrane. In the fore-
wing (fig. 28) the median vein branches from the radius within the wing-stump ;
in the hind-wing the branch is distinctly beyond the cross suture ; radius very dark,
darker than other veins, shaded above and below beyond the middle; median
nearer the cubitus than the radius, dark at the base only, unbranched or with a
variable number of indistinct branches near the apex; if branched, the branches
joining the wing margin at or above the apex; cubitus with 9-12 branches, the
first 4-8 dark and often branched, the remainder lighter, simple or branched once
or twice, all reaching the posterior margin.

Fig. 28. Leucoteymes clavki, sp. n., wings of imago.

Abdomen elongate, very little widened in the middle, bluntly rounded at apex ;
sternites 1-7 with clear spot at each end ; cerci short, basal portion as long as apical.

Measurements: Total length, about 7; head, with jaws, 0-99-1-17 long ; thorax
and abdomen, 5-57 long; head, at and including eyes, 1-2 wide; antennae, 1-7 ;
mandibles, right, 0-61 long, 0-43 wide, left, 0-66 long, 0-37 wide ; prothorax, 0-6 long,
0:99 wide; fore-wing, 12:5 long, 3-66 wide; hind-wing, 11-5 long, 3-8 wide; |
tibia (i) 0-89, (ii) 0-89, (iii) 1-17; abdomen, 1-64 wide. |

Soldiey.—Head bright yellow, palest behind; mandibles dark ferruginous to —
bright yellow; labrum and antennae bright yellow; pro-, meso- and metathorax _
pale yellow, like back of head, remainder of insect whitish. Head, margin of thoracic |
tergites, and abdomen with scattered, moderately strong, reddish hairs.

Head (fig. 29) long, parallel on the sides, rounded behind. Mandibles long, |
moderately stout, of typical form. Labrum long and wide (0-47 long, from anterior
margin of clypeus, by 0-35 wide across the middle), slightly swollen on the sides,
narrowed to the pointed apex. Anteclypeus very short, hardly visible. Post- |
clypeus nearly twice as wide as long, truncate in front. Fontanelle present.
Maxillary palpi with first and second joints half as long as third and fourth ; fifth |
slightly shorter than fourth. Antennae 16- or 17-jointed ; in 16-jointed antennae, |
Ist joint stout, twice as long as 2nd; 3rd shorter and narrower than 4th, shortest ; |
4th as long as 2nd, swollen ; 5th shorter than 4th; 6th as long as 4th ; 7th longer: |

ON SOME AUSTRALIAN TERMITES. 397

in 17-jomted antennae the segmentation is as described in the imago. Gula (see
dotted line in fig. 29) long and narrow, narrowest in the middle, expanded at the
anterior fifth to twice the width at middle, anterior extremity slightly wider than
middle.

Prothorax (fig. 29) a little narrower than head, as wide as metathorax, rounded
and deeply emarginate in front, rounded antero-laterally, sides narrowed to the
rounded and emarginate posterior margin. Meso- and metathorax rounded on the
sides and behind, hind margin slightly emarginate. Legs short and stout, hind
femora very stout; tibial Spurs oa 12;

Abdomen broad, a little wider in the middle than at the base. Styli very long
and slender, apparently always present. Cerci long and slender.

Fig. 29. Leucotermes
clavki, sp. n., head of
soldier.

Measurements: Total length, about 6; head and mandibles, 2:92-3:25; head,
without mandibles, 1-65-2-1; mandibles from base, 1-4; head, 1-08-1-22 wide,
1-03 deep; head, base to fontanelle, 1-17: antennae, 1-7; thorax and abdomen,
3-3-25 long ; prothorax, 0-6-0-66 long, 0-84-0-98 wide : tibia (i) 0-7—-0-8, (ii) 0-61-.
0-65, (iii) 0-98-1-03 ; abdomen, 0-94 wide.

Worker.—Head and thorax pale yellow, a large ferruginous spot at either end of

postclypeus, rest of insect whitish. Head and body sparsely clothed with reddish
hairs.
_ Head rounded behind, widest across the middle. Anteclypeus small, whitish
with a yellow mark on either side, one-third as long as postclypeus, rounded in front.
Postclypeus large, convex, a little more than twice as long as wide. Antennae
14- or 15-jointed.

Prothorax rounded, bent up in front and slightly’ emarginate, sides narrowed to
the posterior margin. Legs short and stout, third femora not greatly enlarged ;
tibial spurs, 3:2: 2.

WESTERN AUSTRALIA: Swan River, Dwellingup, Ludlow (J. Clark).

Type series in author’s collection ; co-types in Mr. J. Clark’s collection.
(5296) 262

398 GERALD F. HILL.

This species is most nearly related to Leucotermes ferox, Frogg. (fig. 30), a co-type
(alate) of which I have examined, but it is easily distinguished by its larger size,
darker colour, much darker and larger wings, larger eyes and fontanelle and short
broad head. The soldier is larger than that of L. ferox. In L. validus, Hill, as in
L. paradoxus, Frogg. (fig. 31), ocelli are wanting in the imago, the fontanelle is minute

and set far back on the head, and the wings are much paler.

Silvestri (1909, figs. 127, 130 and 132) refers certain large-sized soldiers from
Western Australia to L. ferox, Frogg., which I think are most probably referable to
L. clarki, sp.n. The soldiers of all the species mentioned above are very similar and _
in most cases I have failed to find any reliable distinguishing characters ; for this |
reason I have withheld descriptions of apparently distinct species from Queensland,
Victoria and Western Australia until winged forms are available for study.

Pee

Fig. 30. Leucotermes ferox, Bigt® °3l Leucotermes
Frogg., head and prothorax pavadoxus, Frogg., head
of imago ; from a co-type. of imago ; froma co-type.

Biology.

Described from a few imagines, soldiers and workers taken on 13th May from a
small mound about 12 in. high, and from numerous soldiers and workers taken in)

September of the same year.

REFERENCES.
Banks, N. and Snyper, T. E. (1920). A Revision of the Nearctic Termites.=
U.S. National Museum, Bulletin No. 108.

Froccatt, W. W. (1896-7). Australian Termitidae, Parts 1, 2, 3.—Proc. Linnean
Society of New South Wales.

Froceatt, W.W. (1915). White Ants.—Farmers’ Bulletin No. 60 (Second edition) ;
Department of Agriculture, New South Wales.

Fuiier, C. (1920). Studies on the Post-embryonic Development of the Antennae
of Termites——Annals of the Natal Museum, iv, part 2.

ON SOME AUSTRALIAN TERMITES. 399
Hirt, G. F. (1915). Northern Territory Termitidae.——Proc. Linnean Society of
New South Wales, xl, part 1, no. 157.
Hizr, G. F. (1921). The White Ant Pest in North Australia——Commonwealth of
Australia, Institute of Science and Industry, Bulletin No. 21, Melbourne.
Jack, R. L. (1897). Notes on the Meridional Ant Hill of the Cape York
Peninsula.—Proc. Roy. Soc. Qld., xu.

MyOBERG, E. (1920). Results of Dr. E. Mjéberg’s Swedish Scientific Expeditions
to Australia. Isoptera.—Arkiv. f6r Zoologi, xii, no. 15.

Ripeway, R. (1912). Colour Standards and Nomenclature. Washington.
SILVEsTRI, F. (1909). Die Fauna Siidwest-Australiens, Isoptera, ii, no. 17.

THompson, C. B. and SnyDErR, T. E. (1920). The Wingless Reproductive Type of
Termites.— Journal of Morphology, xxxiv, no. 3.

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BuLi. ENT. RESEARCH. VoL. XII. Part 4. PLATE IX,

Fig. 1. Termitarium of Drepanotermes silvestrii, sp. n.

Fig. 2. Termitarium of Drepanotermes silvestril, sp. n.,
showing internal structure.

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BuLLt. ENT. RESEARCH. VoL. XII.’ Part 4. PLATE X

Form of termitarium of Hamitermes perplexus, sp.n., built
on open plains.

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PLATE XI.

ParT 4,

elle

VOL.

Butt. ENT. RESEARCH.

beds 4h) cacc. wale

Oy,

Termitarium of Hamitermes meridionalis, Frogg., near Darwin, Northern

Territory, Australia.

PLATE XII.

Butt. ENT. RESEARCH. VOL. XII. PART 4,

Fig. 1. An unusual type of termitarium of Hamitermes perplexus, sp. Nn.
The track in the foreground has been made by an invading army of ants
(Iridomyrmex).

Fig. 2. Termitarium of H. perplexus invaded by Iridomyrmex, which have
entered at holes marked X. The dark portions have been recently added
by the termites.

401

THE MOSQUITOS OF SOME PORTS OF CHINA AND JAPAN.
By W. A. LAMBORN,
Malaria Bureau, Federated Malay States.

A mosquito survey, having as its object an enquiry into the distribution and
prevalence in certain Far Eastern Ports of Stegomyia fasciata, the known carrier
of yellow fever, was begun, on behalf of the Imperial Bureau of Entomology, by
Dr. A. T. Stanton, whose report was published in the Bulletin of Entomological
Research (Vol. x, pt. 3). As it seemed unlikely that he would be at leisure to
complete the observations by a survey of the ports of China and Japan, as had been
planned, the writer was instructed, in April 1921, by the Government of the Federated
Malay States, at the suggestion of the Imperial Bureau, to complete the enquiry.

The following ports were accordingly visited with this object between April and
June: Fuchow, Shanghai, Nagasaki, Kobe and Yokohama. The object of the
present report is to afford an account of the investigation and of the observations
made.

In a latitude so far north as that in which the enquiry was commenced, namely
26° 8’ 4", it was hardly to be anticipated that a considerable variety of mosquitos
would occur ; nor at the season, early spring, were great numbers of imagos to be
expected. So difficult indeed was it to discover any at all, that it was found necessary
to abandon the attempt to determine the presence of mosquitos by capture and to
concentrate on the investigation of possible breeding places, a method the surer
on account of recent experience, especially in the Malay States, of the breeding
habits of the common CULICIDAE. By breeding, long series of adults were obtained,
for it was found that the larvae of all species were little affected by the shaking they
were subjected to in the course of travel, whether by land or sea. Thus the larvae
obtained in I*uchow, China, afforded imagos fifteen days later in Nagasaki, in Japan ;
imagos were obtained in Yokohama from larvae taken fourteen days previously
in Nagasaki, 853 miles to the south by rail; and imagos emerged on the steamer
in Mid-Pacific, some 2,000 miles from land, from larvae taken eight days previously
at Yokohama.

Fuchow.

Fuchow, the capital of the Fukien Province of China, is situated in latitude 26° 8’ 4”
North and longitude 119° 37’ 7” East on the north side of the Min river at about
35 miles from its mouth. It is 140 miles north of Amoy and 280 south of Hangchow.

The Foreign Settlement is on Nantai, a small isiand formed by the parting and
rejoining of the river and connected with the town on the mainland by an old stone
bridge. The Chinese town itself, with a population estimated at 600,000, consists
of a labyrinth of native buildings, largely one-storied, intersected in every direction
by lanes, usually paved, and so narrow as to allow only foot traffic.

Except for a small hill partly occupied by the Foreign Settlement, which is hemmed
in on all sides by the native quarters, the ‘district is entirely flat and so little above
water-level that with sudden rises of the river, which take place once or twice
annually, the whole town becomes inundated. Rice-fields surround it on every side.

The river is crowded with craft of all descriptions, from large sea-going junks to
stnall two-oar and other boats trading with the interior. The town is not
accessible to ocean-going vessels other than junks, larger ships anchoring about
twelve miles down stream, at Pagoda Anchorage.

402 W. A. LAMBORN.

Trade.—The trade of the port is mostly with other coastal places, for owing to
trade depression, particularly in tea, for which until a few years ago Fuchow was
one of the world’s centres, large ships rarely put in. As showing how great has been
the falling off in trade, it may be mentioned that, whereas in 1893 the total export
of tea was 65,794,894 Ib., it had dwindled to only 6,941,877 lb. by 1918.

Steamers, mostly European-owned, run to and from Hong Kong, while the steamer
trade to Shanghai is largely in the hands of the Chinese. During a part of the year
motor schooners, trading under the Japanese flag, run to and from Formosa, and a
very occasional ship brings kerosene from San Francisco.

Climate.—The seasons correspond roughly with our own, the summers being as
a rule excessively hot, while in winter, when snow may fall, an icy blast sweeps down
over the hundreds of miles of flat treeless country to the north. No data as to rainfall
were available.

Water Supply.—There is no public supply. Water is usually taken from the Min
at low tide and is slightly brackish, but there are a few surface wells in the grave-
yards, the only open spaces in the city. As is usual in native towns without a pipe
supply, water is stored in large earthenware jars, which are also used for collecting
rain-water.

Sanitation and Quarantine
entirely to the demand for excreta for agricultural purposes. Scavengers, usually
women, carrying uncovered wooden buckets, make a house-to-house collection day
by day, and from time to time bale out the contents of earthenware jars disposed
in odd corners for the convenience of passers-by, and vie with each other in seeking
to obtain for sale such casual deposits as may be promiscuously made elsewhere.
The material, after long storage in primitive septic tanks, is dumped on the rice-
fields. The effect of this pollution on mosquito breeding would seem to be a point
well worth determining. Other waste water is either thrown directly into the street,
or, escaping under the walls of the houses into a pit on the outside, forms possible
breeding places. No measures against mosquitos are in force.

A European Medical Officer, under the Chinese Maritime Customs, is stationed
at Pagoda Anchorage ; all ocean-going ships are subject to his inspection and he is
able to enforce quarantine there.

Such hospitals as there are in the town, witha total of about 250 beds, are connected
with the various Missions, which in the past have attempted to deal with any epidemics
that have arisen. During the outbreak of cholera in 1919 which, it is estimated,
resulted in the death of no fewer than 40,000 people, a large number of temporary
hospitals were erected by the American Red Cross.

Mosquitos —The following is a list of the mosquitos obtained : Stegomyia albopicta,
Skuse (= scutellaris, Theo.), Culex fatigans, Wied., C. tr ‘taeniorhynchus, Giles, and
A. hyrcanus, Pall. (= sinensis, Wied.).

Stegomyia albopicta—The larvae of this species teemed in all water-jars left
standing. These seemed to be the only breeding places available to the insect,
which favours artificial breeding places ; for the thrift of the Chinese leads them to
collect all old tins, broken bottles and similar receptacles, such as in the vicinity of
habitations in the Federated Malay States afford abundant opportunities for the
insect.

It was usually possible to determine the species of larvae in particular vessels
by examination of the adults found newly emerged on the inside at almost any hour
of the day. But although long series of larvae were collected from numerous places.
no single specimen of S. fasciata was obtained.

Culex fatigans—The larvae of this species swarmed in the foul-smelling waste
water from houses, and in certain ponds which had once been used for fish cultivation

MOSQUITOS OF SOME PORTS OF CHINA AND JAPAN. 403

but have since served as dumping grounds for all sorts of organic refuse, chiefly of
vegetable origin. They were also found sparingly in several places in ill-smelling
water in pans placed under flower-pots. The species was certainly the dominant
one in the town.

Culex tritaeniorhynchus.—The larvae of this species were found sparingly in the
rice-fields associated with those of A. hyrcanus.

Anopheles hyrcanus.—The larvae of this insect were found in fair abundance on
the outskirts of the town in the rice-fields, then at an early stage of cultivation,
the planting out of the young plants from the nursery beds having only just com-
menced. It was noteworthy that in these beds, longer undisturbed than the open
fields, the larvae were far more numerous than in the newly planted fields, possibly
by reason of the presence of a larger amount of green filamentous algae, on the tuits
of which they were invariably found. The larvae were either green in colour,
or green with white splashes ; or banding was present to a varying degree on the
thorax, 3rd, 5th and 8th segments. They did not present that considerable diversity
both of colour and pattern seen in those of the Malay States. Though they rested
on a green background, it was easy to see them when the sun’s rays were
oblique, but difficult to do so with the sun overhead.

Shanghai.

Shanghai, the great emporium of Central China and its most important free port,
is situated in latitude 31° 14’ North and longitude 121° 29’ East on the left or west bank
of the Huangpu River, about twelve miles above its junction with the Yangtze and
at a distance of sixty miles from the North Saddle light at the river mouth.

Three distinct areas, each having its own municipal council, comprise the port,
namely, the International Settlement, the French Settlement and the Chinese City.
The area within the municipal limits is 83 square miles, and the quinquennial census
of the foreign and Chinese populations residing within the limits and of foreigners
living on the outside roads, which was taken on 10th October 1920, showed a total
of 26,869 foreigners and 1,661,098 Chinese. Among the foreign population the
Japanese were dominant, numbering 10,215 ; the British coming next and numbering
5,341. The figures do not include a vast Chinese population residing outside the
Settlements.

The country round Shanghai, which is devoid of trees, is perfectly flat, stretching
as arich alluvial plain to the west over 45,000 square miles. The area round the town
is little above sea-level, so that it does not present a great variety of possible breeding
places for mosquitos.

Trade.—Shanghai is not only a port of trade, but is a large manufacturing and
industrial centre to which ships come from all quarters of the globe.

Temperature and Rainfall—The annual mean temperature is about 50-9° F.
In July and August it runs up to a mean of about 81°, and in January and February
drops to a mean of 36° and 37°. In winter, snow and ice are occasionally seen,
biting winds sweeping down from the north. A certain amount of rain falls through-
out the year, the annual fall being about 39-97 inches, with a maximum fall of about
five inches in June, July, December and February.

Water Supply—tThe public supply for the International Settlement is drawn
from the Huangpu, below the town ; that for the French Settlement is obtained some
distance above it. It is piped to most of the houses, or is obtainable from standpipes,
so that there are practically no wells.

Sanitation and Quarantine —The Health Department of the International Settle-
ment, under British supervision, is a large one, consisting of a Director with three
Assistant Health Officers, fourteen Sanitary Inspectors, and three Sanitary Overseers,

404 WwW. A. LAMBORN.

with a large native staff. Anti-mosquito measures are thoroughly carried out.
For the purposes of mosquito reduction a special staff of Chinese coolies is organised
from the middle of March until the end of October, and works under the direction
of Inspectors, who make written notes of those places where stagnant water occurs,
such as Chinese gardens, empty houses and defective gullies, such spots subsequently
receiving special attention.

By means of posters and leaflets in the languages commonly spoken an
endeavour is made to instruct the people as to the necessity for mosquito
control, but the difficulties of this are enhanced by the supineness of the Chinese
authorities controlling the part of the town outside the Settlements. Within the
actual limits of these the search for mosquitos was fruitless ; outside it the larvae
of some dominant species were obtained abundantly, and it was said that mosquitos
in the European part of the town are a nuisance, as a rule, only in the late summer,
gradually extending in from the Chinese quarters.

The shipping and quarantine are under the control of the Chinese Maritime
Customs. A medical officer visits ships when necessary and when they come from
infected ports. The quarantine station, where hospital accommodation and fumiga-
tion apparatus are provided, is some miles down stream.

Mosquitos.—The following is a list of the species obtained : Stegomyia albopicta,
Culex fatigans, Culex tritaemorhynchus, C. virgatipes, Edw., and Anopheles hyrcanus.
Three of these species, with Avmigeres ventralis, Walk., are recorded in the report of
the Health Officer of Shanghai for 1920 as having been collected in the course of
examinations made by the staff. The two new records concern C. tritaeniorhynchus
and C. virgatipes.

Stegomyia albopicta.—A search for four days in various parts of the International
and French Settlements entirely failed to bring to light any larvae of Stegomyza.
On the fifth day access was obtained, through the kind intervention of Mr. E. Kilner,
Chief Sanitary Inspector, to a Chinese-owned greenhouse, standing by itself
and not at that season artificially heated, where in the water under five out of eleven
little fern-covered rockeries kept in pans the larvae of this species were for the first
time obtained. They were subsequently found in one other similar situation, but a
further search for them elsewhere was entirely fruitless. No other species of
Stegomyra was obtained. The scarcity of this insect at that time may well have been
due to inclemency of season, which was said to have been unusually cold throughout,
the mean daily temperature during the stay of five days in Shanghai being 61°,
a bitter northerly wind blowing.

Culex fatigans.—The larvae of this species were found in open drains from houses
and in the trenches draining vegetable plots.

Culex virgatipes.—This species was found in the same breeding places with
C. fatigans, but in greater abundance.

Culex tritaeniorhynchus.—A single female example of this species, of very large
size, was bred from a pupa found in a small swamp associated with those of
A. hyrcanus.

Anopheles hyrcanus.—The larvae of this species were obtained in fair abundance
n foul and stagnant water in swamps on the outskirts.

Nagasaki.

Nagasaki is situated in latitude 32° 45’ North and longitude 129° 52’ East, at
the head of an inlet some three miles long at the western extremity of the island
of Kyushu, which is about 469 miles from Shanghai. Being the most southerly
of the ports of Japan it is the first port of call of the eastward steamers from India,
the South Seas, China and the Philippines. It is largely built at the foot of hills

MOSQUITOS OF SOME PORTS OF CHINA AND JAPAN. 405

forming a basin, but the houses extend up into the valleys to an elevation of about
400 ft. It has a population estimated at 176,480, and, in point of size, ranks next
after Kobe in Western Japan.

Trade.—It is noted as a coaling station, the coal being obtained chiefly from
Takashima, an islet eight miles south-east of the entrance ‘to the harbour, and in
lesser quantities from two adjacent islets.

There are no large industrial concerns in the south of Japan other than the
shipbuilding vards at N agasaki, and the exports are therefore small.

Temperature and Ratnfall—rThe climate is mild and equable, the mean annual
temperature being from 60° to 63° F., with extremes of 40° to 45° in winter, and
75° to 85° in summer. Snow and frost are unknown.

Water Supply.—-Mountain torrents supply the outlyiz ing parts of the town with
water ; the supply to the centre is pipe-borne from reservoirs in the hills near by.

Sanitation and Quarantine.—A quarantine station, with shore hospital accommoda-
tion and apparatus for the fumigation of ships, exists at a distance of a couple of
miles to seaward of the harbour, and similar provision is made at al! the other large
ports of Japan.

Mosquitos.—A greater number of species than was met with in Fuchow and
Shanghai was to be expected in Nagasak!, by reason of the more genial climate and
the greater variety of breeding places afforded by the natural features of the country.
The following species were obtained : Stegomvia albopicta, A édes (Finlaya) togot,
Theo., Aédes (Finlaya) japonicus, Theo., Culex fatigans, C. hayashi, C. tritaenio-
rhynchus, Armigeres obturbans, Walk., Lutzia vorax, Edw., Anopheles hyrcanus,
A. lindesayi, Giles, and Anopheles punctibasis, Edw.

Stegomyia albopicta.—The larvae of this species were fairly abundant in artificial
breeding places, usually in shaded situations and in pure culture. They were
obtained in rain-water collected in earthenware jars, in saucers beneath flower-
pots, and in the cement or stone basins beneath the little fern-covered rockeries which
are so commonly seen in Japanese gardens. An oblong granite trough, shaded
by the eaves of a house in the main street, afforded a vast number of larvae, and
similar troughs, set beneath a roof of thatch in the court-yards of temples and
containing water for worshippers to rinse their mouths and wash their hands, pre-
paratory to entering the sacred edifices, almost invariably afforded larvae. Their
security in such situations is to be explained largely by the wariness exhibited, the
larvae all dropping to the bottom on the slightest disturbance of the water, and by
there being no means of draining the water from the bottom of the receptacles, w hich
are never completely empty. A little bowl, scooped in the top of a granite pillar
placed in front of a stone image of Buddha in a small roofed wayside shrine, afforded
larvae, which were also found there in the water in sections of bamboos placed for
the reception of votive offerings of flowers. No specimens of S. fasciata were
obtained.

A édes (Finlaya) togoi.—-The larvae of this species were first obtained in enormous
numbers in the centre of the town, on the hillside in a large uncovered cement tank
holding some hundreds of gallons and used for the storage of rain-water. They
were invariably found in the: water-containing troughs cut in the solid stones forming
the pedestals of the monuments in cemeteries, and in the granite vases, often of very
large size, placed for ornamental purposes in gardens. Cups hollowed in boulders
placed near the doors of houses, and containing water used for ablution purposes,
often contained these larvae when in the open, and they were commonly found in
water-containing hollows in rocks in partly dried-up beds of streams running down
from the mountains through the town

406 W. A. LAMBORN.

Aédes (Finlaya) japonicus.—This, unfortunately, was not recognised as a distinct
species, until Mr. F. W. Edwards had made the determinations. It was found less
abundantly than A édes togot, though breeding in similar places, and, it is believed,
mixed with this species.

Culex fatigans —The larvae of this species were found abundantly in foul drains
and cesspits. Such was the cleanliness of the town, in spite of the absence of any
water-borne sewage system, that these breeding places were not discovered at all
within its precincts. Open drains were found only in the vicinity on the country-
side, usually about cattle-sheds and rarely near houses. Pits, open to the sky, either
dug in the ground or cut in the rock, were dotted about the fields, and served as
septic tanks into which collections of excreta were dumped, the contents being
finally withdrawn for the purposes of agriculture. In such pits, whether containing
comparatively fresh or thoroughly decomposed material, the larvae invariably
occurred in enormous numbers, in the former case in pure culture.

Culex hayashi.—The larvae of this species were found in great abundance
associated with those of (Anopheles punctibasis) in a small natural pond, almost
dried up and completely shaded at all hours by trees.

Culex tritaeniorhynchus.—The larvae of this species occurred, as elsewhere, in
rice-fields.

Armigeres obturbans.—A few imagos of this mosquito were obtained in houses.
Their breeding places were not discov ered.

Lutzia vorax.—The larvae of this species invariably occurred in some abundance
in cesspits, though only when the contents were so thoroughly decomposed that
algal growth had become possible. In such pits the larvae of C. fatigans were z
less abundant then elsewhere, the reason for which became apparent when about <
dozen larvae of the Lutzia were collected into a bottle. They attacked each athe
with such ferocity that within an hour only two or three survived, and they fed freely
on the larvae of fatigans on which a long series were bred to maturity. The pupae
of this Culex however seemed to enjoy entire immunity from attack, possibly because
at the surface they are not so readily seized by such assailants.

Anopheles hyrcanus.—The larvae occurred in great abundance in the rice-fields
when at an advanced stage of cultivation, associated, as in Fuchow, with those of
C. tritaeniorhynchus. As in Fuchow, thoroughly decomposed manure in a liquid
state is poured on the fields, when they are first flooded ready for planting out the rice,
but in Nagasaki a great abundance and variety of waste vegetable matter, such as
cut grass, potato haulms, turnip tops, etc., are also thrown in and allowed to
decompose before cultivation is commenced, the larvae being seemingly unaffected
by the richness of the water in organic matter. In Nagasaki, as elsewhere in Japan,
the larvae were similar in colour and pattern to those in the Chinese ports referred to.

Anopheles lindesay1.—Nine larvae, only, of this species were obtained, in clear
chilly water bubbling up from a spring in the hillside, a habitat similar to that in which,
as Dr. Hacker has written, he found them in India.

Anopheles punctibasis.—The larvae of this species were obtained in some
abundance in the same muddy pool with those of C. hayashi. The insect has
recently been described by Mr. F. W. Edwards (Bull. Ent. Res. xii. pt. 3, p. 274,
Nov. 1921).

Kobe.

Kobe, situated in latitude 34° 41’ North and longitude 135° 11’ East on the
Inland Sea of Japan, has recently superseded Yokohama as the principal port, the
tonnage of vessels arriving and clearing amounting in 1919 to 36,100,000, for it is
accessible to the largest steamers. The town is situated on a strip of land from a

MOSQUITOS OF SOME PORTS OF CHINA AND JAPAN. 407

half to one mile wide, between hills at the back and the sea, and so has a very long
sea frontage. The population in 1919 was 588,124, and the town is distant only 20
miles from Osaka, also on the coast, with a population of 1,400,000. There is ample
communication between the towns by steamers, trains and electric trams.

Trvade.—-The main trade of the Empire centres in Kobe, steamiers from all quarters
of the globe berthing there, after having, as a rule, first visited either Nagasaki or
Yokohama. <

Temperature and Rainfall—These are practically those of Yokohama.

Water Supply.— A piped supply is everywhere laid on from reservoirs in the
mountains at the back of the town.

Mosquitos—The following species were obtained: S. albopicta, A. (Ftnlaya)
togot, A. (Finlaya) japonicus, C. tritaeniorhynchus, C. fatigans and A. hyrcanus, the
larvae occurring in places similar to those in which they were found in Nagasaki.
A species regarded by Mr. F. W. Edwards as C. pipiens, L., though the male genitalia
are not quite identical with those of European specimens, was here obtained for
the first time, breeding in similar places to C. fatigans, if not mixed with it.

There seemed also to be a marked reduction in A. japonicus, for one specimen
only was obtained, though A. fogoz occurred in great abundance.

Yokohama.

Yokohama is situated in latitude 36° 26’ North and longitude 139° 38’ East on
Tokyo Bay, 18 miles south-west of Tokyo. It stands on a plain shut in by hills on
either side. It is the sixth largest city in the Empire, with a population of 400,000
Japanese and 10,000 foreigners, who mostly reside in one quarter of the city.

Trade.—Its natural excellence as a harbour has made it one of the two greatest
ports in the country, its imports and exports being second only to those of Kobe.
All the steamer lines from China, India and Europe converge at Yokohama, and it
is the first port of call for Trans-Pacific liners from the United States and Canada.
It is in direct communication with ports of the west coast of Mexico and Central
America, the ships of one of the large passenger lines, coming by way of the Panama
Canal, taking from 23 to 27 days in making the journey from Panama to Yokohama,
putting in at San Francisco en route.

Temperature and Rainfall—The seasons correspond to our own, the mean tem-
perature in the spring (March to May) being 55° F., in the summer (June to August)
73-9°, in the autumn (September to November) 60-6°, and in the winter (December
to February) 39°. The cold in winter is often severely felt owing to northerly winds.
The rainfall is about 70 inches annually. There is a heavy rainfall during the winter
and early spring months, and after an intermission of a couple of months, mild rains
fall with some constancy in June. The latter rains, coming at a season when with
rising temperature mosquito activity begins to be felt, serve to keep possible breeding
places constantly full of water, a factor of importance as favouring at that season
the increase of mosquitos. At Kobe, and in the south, these rains are less constant.

Water Supply.—-A piped supply serves the whole city, but the collection of rain-
water in unscreened vats and barrels is general, there being a demand for it for the
dyeing of cloth.

Mosquttos.-—The following were obtained: Stegomyra albopicta, Aédes (Finlava
. . . . 5 . 6 7 4 i
togot, Culex pipiens, C. orientalis, Edw., and Anopheles hyrcanus.

Stegomvia albopicta.—The breeding places of this species were of the same type
as in Nagasaki, though the larvae were obtained less freely than in the south. They

408 w. A. LAMBORN

were found, for example, in a hollow tree in one of the main streets, but were especially
abundant in the cemetery of the Foreign Settlement, where plentiful breeding places
were provided for them in the jam-jars, pickle-bottles, and similar inexpensive
receptacles that had once contained those floral tributes, which, in accordance with
pious custom, are at lengthening intervals placed upon the graves—auntil the decay
of interest in the departed gives the female mosquito the reversion. The specimens
here obtained were so diminutive as to make their recognition, which is usually so
easy by reason of the very characteristic markings, difficult without the aid of a
lens. Specimens taken in Fuchow and Shanghai were rather larger, but did not
come up to the size of those ordinarily taken in the Malay States. The gradual
diminution in size may well be due to conditions becoming more unfavourable to
the species further north, for it was found repeatedly in the Malay States that poorness
of the food supply resulted in the production of dwarfed imagos. The ability to
attain maturity at all under such conditions is doubtless one of the reasons why this
species and S. fasciata are so widely spread. All attempts to obtain in the Malay
States dwarfed Anophelines by semi-starvation or by rearing them in unsuitable
media failed : the larvae either attained a maximum growth and in due course afforded
imagos, or else perished after a prolonged period of larval existence.

A édes (Finlaya) togot.—-This species was found in the various types of breeding
places already alluded to. It was especially abundant in certain quarters in butts
of rain-water, and large numbers were obtained in the yards of stonemasons, in the
cavities of various granite receptacles, such as mortars, bowls and troughs of all
sorts, shapes and sizes, ornamental vases, etc. At a particular temple, at the main
doorway, the pillars on either side rested in huge iron tubs, holding gallons of water.
Examination for larvae in one of these, in which there were three golden carp, was
negative ; in the other, in which there were no fish, larvae swarmed. It had already
been remarked that in none of the little ornamental ponds, invariably found in gardens
of any size, were larvae found when goldfish were present. It was quite usual to
find a few of these fish in surface wells and barrels of water, in which, again, no larvae
were ever obtainable. While these larvae were invariably found in open situations,
their near relation, S. albopicta, was found breeding, with very few exceptions, in
thoroughly sheltered spots. The larvae of Aédes japonicus were not obtained here
at all.

Culex orientalis —The larvae of this species were found associated with those of
Anopheles hyrcanus, having seemingly replaced those of C. tritaeniorhynchus, which
were abundant in the south.

Culex pipiens.—This again had entirely replaced C. fatigans,and was found breeding
in great abundance in similar drains and ditches.

Anopheles hyrcanus.—This was found in abundance, breeding, as usual, in rice-
fields.

Conclusions.

Confirmation of the opinion expressed by Dr. Stanton in his report that “a
survey of the ports of China and Japan would show that the conditions are unfavour-
able for the propagation of Stegomyia fasciata, even in the warmer months ”’ was,
therefore, obtained in regard to the places visited. In view of a record (Theobald,
Monograph of the Culicidae, i, 1901, p. 293) of the capture of a single specimen of
the insect in Tokyo (C. H. B. Wood, 3.viii.1899), a three-days’ search for it was made
in the suburbs there, especially on the seaward side. None were obtained, so that
one may well conclude that, whatever the history of this particular specimen, the
species has failed to establish itself there. There are recent records, however, of its
presence in Kowloon, in latitude 22° 12’ North on the mainland opposite Hong Kong,
though sparingly in the latter place (Stegomyia Survey in Hong Kong, Bull. Ent.
Res., vi, 1915, p.67), and in Formosa (Secrete, 1917, ‘‘ Notes of Mosquitos of Formosa.”

MOSQUITOS OF SOME PORTS OF CHINA AND JAPAN. 409

Abstract in China Med. Journal, Shanghai) in latitude 23° 5’ North, and so occasional
specimens may find their way north, though as a factor in the spread of yellow fever,
they would appear to be negligible.

The possibility of the establishment of yellow fever in the ports visited would
seem to hinge on the potentiality of Stegomyta albopicta, and possibly other very near
relatives, such as A édes togot and A. japonicus, to act as carriers of the disease. Mr.
Kilner stated that in Shanghai these mosquitos are not seen during the colder months
of the year, and this probably holds good of Fuchow, where bitter cold is also experi-
enced. In Japan, which feels the benefit of the North Equatorial Ocean current,
the seasons, especially in Nagasaki, are milder, and it would seem probable that
here, with a mid-winter temperature no lower than 43° 5’, imagos may be active all
the year round.

The work was greatly facilitated in Fuchow by the kind and ready help of Dr.
Cheah and of Mr. H. S. Brand, the Secretary of the local Chamber of Commerce.
I must acknowledge, further, the assistance received in Shanghai from Dr. C. N. Davis,
the Acting Director of the Department of Public Health, and from Mr. E. Kilner,
Chief Sanitary Inspector, and express my indebtedness to Mr. F. W. Edwards for
the determinations of the mosquitos obtained.

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411

NOTES ON THE SPECIES OF THE GENUS MUSCA, LINNAEUS—PART I.
By Major W. S. Patron, M.B., F.E.S., I.M.S. (retired),
Lecturer on Entomology and Parasitology, Edinburgh University.

Of the many insects which we now know to be dangerous menaces to the health
of man, the house-fly, Musca domestica, has few (if any) equals, but the facts relating
to its germ-transmitting capacity are unfortunately still very incomplete. In the
case of those Anopheline mosquitos which transmit the parasites of malaria, and
the tsetse-flies, which infect man and other animals with trypanosomes, we are in a
position to estimate the inefficiency and loss of life directly attributable to them, for
in each case we know most of the species directly responsible for the dissemination
of these parasites. But in the case of the house-fly we have no conclusive proof
that Bacillus tvphosus, for instance, is carried by one, or more than one, species of
Musca. Further, by the time the cases are diagnosed, the infected flies have either
died or disappeared, and it is rarely possible to trace the infection to them. It is
true, however, that many observers have recorded the finding of pathogenic bacteria
in, and on, species of Musca, but the final proof of the identity of these bacteria has
invariably been faulty. Yet in spite of these discrepancies there is no doubt, from
everyday practical observation the world over, that one, and more than one, species
of Musca regularly carry the bacilli of the enteric fever, cholera and dysentery groups,
as well as many other pathogenic organisms, especially the virus of trachoma, from
infected material and deposit them on food and the human body. In investigating
outbreaks of infectious diseases the house-fly should always be considered as a possible
vector of the causal organisms.

The fact that the CULICIDAE contain among their members many serious pests
has led to an intensive study of the species of the world, resulting in numerous
valuable monographs and papers describing the species, and every medical man is
now able to determine with certainty almost any species for himself. So also in the
case of the tsetse-flies we have Austen’s complete handbook available for the deter-
mination of the species ; and similarly with many other insect pests. This systematic
work on these important insects is very necessary, for it facilitates further
biological studies, and, more particularly, accurate information on their relation
to disease-causing germs.

But when we come to consider the all-important house-fly, Musca domestica
and its allies, we find that although there are many valuable papers and books dealing
with this species from the disease standpoint, there is no systematic work of any value
which would enable the sanitary officer in the tropics to determine with accuracy
the species which may call for enquiry during the course of his work. He cannot
help but call all the species of Musca by the specific name domestica, for he believes
that a house-fly with a striped thorax is Musca domestica all the world over. And
there is every reason that he should do so, for in such an authoritative work as that
on “‘ The House Fly, Musca domestica, L.”’ by the late Dr. Gordon Hewitt, we have
a definite statement that the house-fly found throughout the world, and especially
in tropical countries, is Musca domestica, L., and this statement has been accepted
and copied by all subsequent writers on the subject. I propose to question it. That
both the sexes of one of the species of \wsca found in the tropics are very similar to
Musca domestica is quite true, but when we come to compare the male with the male

(5296) 2H

412 MAJOR W. S. PATTON.

of domestica, we find that the front of the former is at least half as wide as that of the
latter. Were this only the case in a few specimens, I should be prepared to accept
Dr. Hewitt’s statement regarding the world-wide distribution of Musca domestica,
but after examining hundr eds af specimens of males of this tropical species,
and comparing them with typical males of domestica caught in Great Britain,
which I assume are conspecific with the Linnaean species, I find that in not a single
specimen from India which I have examined is the front as wide as that of the male
domestica. It is true that the width of the front of this form varies to a small
extent, and so does that of the typical male domestica, but on the one hand I have
yet to see a specimen of domestica from Great Britain with a front as narrow as that
of this tropical form, and on the other hand a specimen of this tropical form with
a front as wide as that of the typical domestica.

I am quite prepared to find a typical male of domestica taken at any of the tropical
ports visited by ships from Europe, for everyone who has travelled in the East knows
that large numbers of domestica are regularly carried to such ports, and leave the
ships on their arrival at their destinations. In this way it is possible Musca domestica
has been carried to the ports of the world, and has now become established there.
But I must admit that the examination of the specimens in the National
collection from the localities mentioned by Dr. Hewitt have not convinced me that
this is the case.

It is true that Major Austen and other authorities have drawn attention to this
narrow-fronted male house-fly and have determined it as domestica, L. But when
we find a form exhibiting a character which is constant, I think we are justified
in regarding it as distinct. And the width of the front in the males of the species
of this genus is a constant character, and one of the most reliable for purposes of
determination. In using this character it is very necessary to point out that great
care must be taken in noting whether the front has collapsed, which it often does
in the case of specimens pinned too soon after hatching and before the chitin has
hardened. It should be remembered that just below the front there is an opening
through which the ptilinum is protruded and later invaginated. If the insect is
pinned before the edges of the ptilinal opening have sufficiently hardened, the whole
front is very apt to collapse inwards ; it then appears to be very much narrower
than it really is, and it is only when such a specimen is macerated in caustic potash
that the exact width of the front can be determined with accuracy. Further it is
important to note that in comparing the front of one specimen with another it is
very necessary to take into account their relative sizes. It is always better to com-
pare the width of the front with the entire width of the head, or with that of one
eye, rather than give it in linear measurement.

It may be thought that the question regarding the true identity of this tropical
and subtropical house-fly is only of academic interest, and not worth further investi-
gation, but I may point out that the biological connection of particular species of
insects with certain disease-causing bacteria is now regarded as of a much more
specific nature than has been thought to be the case hitherto. As a good example,
I may draw attention to the case of the tropical rat-flea, Xenopsvlla cheopis, which
is the invertebrate host of Bactllus pestis. Until recently it was thought that there
was only one species of the genus Xenopsylla found on rats in India, but a more careful
study of rat-fleas has enabled Mr. Charles Rothschild to separate the Indian cheopts
into three distinct species, one of which, X. astia, has undoubtedly been confused
with cheopis in the past. It is thought by Cragg and Hurst that the absence of
plague in certain areas is due to the larger percentage of astia found on rats as
compared with the numbers of cheopis. If this is the case, then we have a clear
indication that aséza is not a good host for the extracorporeal life of Bacillus pestis.
In the same way there may be a much more intimate connection between certain
disease-causing bacteria and a particular species of Musca rather than with the others

SPECIES OF THE GENUS MUSCA, LINNAEUS. 413

which may feed side by side withit. Tor these reasons, then, I consider it is important
to determine the true identity of this tropical house-fly. But in order to settle this
question beyond any further doubt, it will be necessary to examine and compare
the external genitalia of both sexes with that of Musca domestica, and to do this a
large number of specimens from many parts of the world will be required, and this
is one of the reasons I have raised this point, in the hope that this material will be
collected. I have already begun the study of the external genitalia of the Indian
species in order to compare it with that of domestica.

But the genus Musca contains another important group of species which have a
direct bearing on tbe health of valuable stock animals, and which have in the past
been confused with Musca domestica. The species IJ refer to, largely if not entirely,
feed on blood and serous and pustular discharges, and are exclusively outdoor insects.
Anyone who has watched a milch cow being tormented by hundr reds of specimens
of Musca autumnalis can hardly doubt that, apart from the possibility of their
carrying disease germs, they must reduce the milk-producing capacity of the
animal simply through wearing it out, as it constantly tries to drive them away.
That flies do in this way so worry a cow as very materially to reduce the quantity
of milk it produces, has been proved beyond any doubt by observers in America.
These species of Musca regularly follow the true biting flies, sucking up any
remains of blood that exudes when the latter withdraw their proboscides. They also
regularly feed on the’ discharge from sores, cuts, eyes and the nose. We know
nothing of the germ-carrying capacity of these species, and until all have been

systematically studied, and veterinary officers and others are able to recognise them,
we never shall.

During the last 15 years I have almost continuously studied all the stages of
many of the species of this genus, and observed their habits i in the field, for i am a
strong believer in combining systematic studies with field observations. I have
no doubt whatever that such field work sives one a much truer picture of the individual
species than mere microscopic study of the dried insect. Field observations, in
conjunction with studies of the br eeding habits and the larvae, as well as microscopic
studies of the adults, are the only means that I know of for coming to a final conclusion
regarding some of the species of this genus. But as an isolated observer in the
tropics, without access to the scattered literature on the subject, I have been severely
handicapped in my systematic studies of the species. I have in the first instance
had to depend on others for determinations of the species, and also on the inaccurate
references to them in the general literature of the subject. As a result many errors

have crept into my papers, and for the benefit of those who may use them I will
draw attention to them in these notes.

Recently I had the first opportunity of studying the specimens in the National
Collection at the British Museum, comparing them with the Indian species, studying
many types, and lastly of consulting the complete literature on the subject. And
I would like to take this opportunity of thanking Major E. E. Austen, D.S.O., for
all the facilities and help he gave me. I have no hesitation in saving that were it
not for his extensive knowledge of the subject, which he freely placed at my disposal,
I could never have done as much as I was able to in the short time available.
My own collection, together with the splendidly arranged and rich National Collection,
has enabled me to unravel almost all the important synonymy. I have critically
examined all Walker’s types, as well as Bigot’s, which Mr. Collin kindly lent me
for the purpose, and for which I wish to take this opportunity of thanking him.
Although I fully realise that much has vet to be worked out before it will be possible
to be certain of the true identity of some of the species of the older writers, I consider
that my study of this extensive material is sufficiently important to call for immediate
publication.

(5296) 2H2

414 MAJOR W. S. PATTON.

But my main reason for publishing these notes is to draw the attention of medical
and veterinary officers, and others, to the importance of collecting more material,
for in spite of the many thousands of specimens which I have examined, I have no
hesitation in saying that my revision of the genus will not be as complete as I should
like it to be. I would, therefore, earnestly appeal to all who have opportunities
of collecting species of A/usca to send them to me at the Zoological Department,
Edinburgh ‘University, or to Dr. Guy A. K. Marshall, C.M.G., Director, Imperial
Bureau of Entomology, British Museum (Natural History), Cromwell Road, when
sending other entomological material ; I shall be glad to exchange any Indian species
for others.

As many of the types of the older writers, such as those of Robineau-Desvoidy
and Macquart, are either lost or in very bad preservation, it is most important to
try to secure specimens from the localities from which their types were obtained,
and the localities will be mentioned in these notes. In order to facilitate the collec-
tion of this material, it may be useful to give some notes on the species as a whole,
and also to indicate the special lines along which the collecting should be carried out.

For practical purposes, the species of the genus Musca fall into two natural
groups as follows: (1) The house-fly group, and (2) the wild species group.

‘1) The House-fly Group.

In this group are included all those species which are cosmopolitan in habit, but
are mainly found in and around human dwellings, on food in bazaars, about butcheries,
offal, rubbish of all kinds and night-soil trenches. The four important species are
Musca domestica, Musca domestica (atypical), Musca nebulo and Musca humilis.
The first is the conmmon [uropean house-fly and the type of the genus, and AZ. domestica
(atypical), M. nebulo and M. humilis are the important tropical and subtropical
species. Specimens of this tvpe of house-fly are urgently wanted from as many
localities as possible. AZusca humilis is a very characteristically marked species,
and can be easily recognised ; but when we come to Musca doniestica (atypical) and
Musca nebulo it is not possible at present to give a definite opinion regarding them.
A series of specimens from the different ports as well as specimens from inland towns
must first be available for microscopic study. Specimens taken 7m copula would
be most useful. Larvae and flies hatched from them are the best material for
comparative study. If the observer has not the time to pin the specimens they will
do equally well if they are placed in match-boxes without any cotton-wool. Such
specimens are quite suitable for the study of the external genitalia.

In collecting these species it is important to note exactly where the specimens
were caught, whether on food, indoors, or in bazaars; and in this connection it
should be noted that this type of Musca may be seen on cattle and horses in company
with specimens of those of the next group. There should be no difficulty in collecting
the larvae of the house-fly in any locality. When mature they should be placed in
some earth in a cigarette tin, the lid of which is perforated with small holes, and
allowed to pupate. At the same time some specimens should be preserved by dropping
them into boiling water in a test-tube, and then placing them in a tube containing
70-80 per cent. alcohol. As soon as those in the tin have pupated, the puparia
should be collected and placed in dry tubes, and 36 hours after the flies have hatched
out they should be pinned along with their puparia; the latter are most useful in
correctly associating the larvae with their respective adults. A good series of males
and females collected in this way would be most valuable for comparative study.

(2) The Wild Spectes Group.

Some years ago I drew attention to the peculiar habit of Musca patton1, Austen,
which feeds on the blood that exudes from the punctures made by biting flies,
and on the serous fluid which exudes from cuts, sores and particularly from cowpox

SPECIES OF THE GENUS MUSCA, LINNAEUS. 415

vaccine scarifications, on the bellies of calves. Since then, in collaboration with
Major Cragg, I.M.S., I have described several other Indian species that have a similar
habit. Looked at with the naked eye, the species of this group closely resemble Musca
domestica. I have never seen a single specimen of any of these species caught in the
bazaars in India, and with the single exception of AZusca humulis, they are funnel only
on animals, sitting about on plants, and on cow-dung. They are all necessarily
intermittent feeders, and flit about from one animal to another, never staying long
at one spot ; and herein lies their importance as the possible carriers of trypanosomes
such as those which cause nagana and surra. I would, therefore, particularly like
to draw the attention of veterinary officers practising in the tropics to these
haematophagous species of the genus Musca. They abound in all tropical countries,
and specimens can always be caught on animals, as well as on dead game.

All the Indian species I have studied breed in cow-dung, either when it is dropped
in the fields, or when piled up in heaps around cowsheds, etc. The larvae of all
the species in any given locality can be easily collected by scooping up patches of
cow-dung about 24 hours old, and placing it on earth in a tin tray with sides up
which the larvae cannot crawl. There is no need to disturb the dung, but the tray
should be placed in the sun every day, and the earth at the sides of the tray
regularly examined for pupating larvae. Such larvae are of a characteristic creamy
white colour, and do not contain any food. They should be collected, some pre-
served as noted above, and the remainder allowed to pupate in earth in a cigarette
tin. The puparia should be dealt with as noted above, always remembering to pin
the puparium of each specimen under it. The mature larvae of most of the species
leave the dung to pupate, but others may stay just under it, so that it is necessary
towards the end of the observation to turn over the dung and examine it for puparia.
The puparia of several of these species are almost white, while*others are of a greyish
colour.

THE ORIENTAL SPECIES.

In addition to the specimens from this region in the National Collection, I have

e-examined my own extensive collection consisting of about 2,500 specimens, the
pescrity of which have been bred from larvae, most of the species being represented .
by long series in perfect preservation. J am able then to note the variations i in external
characters, a point of considerable importance in determining an atypical specimen ;
and anyone who has examined many species of this genus will have noted the great
difficulty often experienced in giving a name to an odd specimen which is in some
respects atypical, or greasy. I also have a long series of bazaar flies collected from
all the larger towns in North India.

In these notes I propose recording my preliminary studies of the Oriental species,
but it should be understood that as more material becomes available for macroscopic
and microscopic study, it may be necessary to revise any previous conclusion, so
that these notes must not be considered to represent the final revision of the Oriental
species, but rather a preliminary contribution in that direction. In collaboration
with Mr. Senior-White, | am now preparing a paper in which all the Oriental species
will be described and illustrated ; we hope to be able to include in it a simple yet
accurate key for the determination of these species.

1. Musea nebulo, Fabricius.

Synonyms: Musca determinata, Walker.
? Musca multispina, Awati.

In 1910, Major Austen determined the common Madras house-fly for me as Musca
nebulo, and since then I have always referred to this species under that name.
fabricius, in his short and wholly inadequate description of this species, states that
the type, a female, is from “ India Orientalis,’’ and was given him by Professor

416 © MAJOR W. S. PATTON.

P. C. Abildgaard. In order to discover the exact locality from which the type was
obtained, Major Austen kindly searched the literature to see whether this Professor
P. C. Abildgaard was ever in India, but beyond the fact that he was a Veterinary
Surgeon, and interested in natural history, there is nothing to show whether he had
travelled in India. The exact locality from which the type of Musca nebulo came
must then, in the meantime, remain doubtful. I think, however, that it can be safely
assumed that it was obtained from some large Indian town. Wiedemann, who
evidently examined the type, does not give any information on this pot. His
somewhat fuller description, however, fits this common Indian house-fly very well.
Short of examining the type I see no reason to doubt Major Austen’s determination
of its identity, and consider it is best to retain this name for it.

I have carefully, and repeatedly, examined the type of Walker’s determinata,
a female in bad preservation, from the East Indies (not the West Indies as
erroneously stated in my paper on the Mesopotamian house-flies), and after com-
paring it with typical specimens of the Madras house-fly (M/Z. nebulo), have come to
the conclusion that it is identical with it. There seems to be no means of discovering
from what part of India the type of determinata was obtained, but the town of
Calcutta seems to be the most probable locality. In my paper on the Mesopotamian
house-flies referred to above, the species there described as Musca determinata,
Walker, is not conspecific with Walker’s type, but is the next species recorded in these
notes.

Although I have not had an opportunity of examining the type of Awati’s Musca
multispina, 1 have little doubt from his description of the female that it is Musca
nebulo.

I have examined many hundreds of specimens of nebulo from many localities in
India, Burma, Assam and Ceylon, and find that though it varies to some extent in
colour and markings, it is a very distinct species and cannot be confused with either
Musca domestica or Musca domestica (atypical) noted below. The majority of the
specimens when in good preservation have white cheeks, a grey thorax and yellowish
abdomen, with silvery tessellation. In the male the frons is narrow, about as broad

_as the width of the third antennal segment (antero-posteriorly), and although in some
specimens it is a trifle narrower, and in others a little wider, I have not seen a single
specimen whose frons in any way approaches the width of the frons of the male
Musca domestica.

I shall be glad to receive any specimens of this species from any area outside
India, so as to compare them with those found in various parts of India.

2. Musca domestica, L. (atypical).

Synonyms: Musca biseta, Hough.
Musca divaricata, Awati.
Musca determinata, Patton (nec Walker).

This species, which for the lack of a better name I propose in the meantime calling
Musca domestica (atypical), is the one which comes very near the true European
Musca domestica, so far as external characters are concerned ; it is one of the other
important house-flies of India. Both sexes have been described by Awati under
the name divaricata, and this observer points out that it closely resembles nebulo
(multispina, Awati), but can be distinguished by noting that it is a much yellower
fly than nebulo, and the abdomen is not nearly so silvery. The frons of the male
of this species is about as wide as that of the male mebulo, and is from a quarter to
one-fifth the width of the eye. I have not seen a single specimen of the male of this
species with a frons as wide as in the male of domestica. I consider it is there-
fore distinct on this character alone. It is, however, not possible at present to give

SPECIES OF THE GENUS MUSCA, LINNAEUS. 417

it a name, and it will be necessary to compare the external genitalia of both sexes
with those of domestica, L., as well as with those of specimens from other tropical
countries. I believe the following will prove to be this atypical form of domestica :—
Musca sanctae-helenae, Macquart, Saint Helena.
Musca lateralis, Macquart, Mauritius.
Musca basilaris, Macquart, Brazil, Mexico.
Musca frontalis, Macquart, Algeria.
Musca analis, Macquart, Chili.
Musca consanguinea, Rondani, America, Mexico.
. Musca senegalensis, Macquart, Senegal.
Musca flavinervis, Thomson, Ross’s Island.
Musca antiquissima, Walker, Australia.
10. Musca calleva, Walker, South Africa.
11. Musca vicaria, Walker, New Zealand.
I have examined Bigot’s types of Musca pampasiana (Buenos Ayres), Musca
flavifactes (New Caledonia), and Musca atrifrons (Cuba), and consider that they are
all specimens of this form. There seems very little hope of getting any further

CON AU WN *

‘light on Macquart’s species, as Mr. Collin tells me that his types are in bad preservation.

When examining these atypical forms of domestica in the National Collection,
Major Austen drew my attention to its occurrence in Greece in company with the
typical domestica, but exactly where this species first makes its appearance in the
Palaearctic Region, I am not at present able to say. I hope those who have oppor-
tunities of examining and collecting specimens of Musca from E urope will be on the
look-out for this narrow-fronted male domestica, and send me specimens.

I have no doubt that Hough’s Musca biseta is this species ; it is common in Aden
and most probably in Somaliland.

3. Musca humilis, Wiedemann.

Synonyms: Musca primitiva, Walker.
Musca conducens, Walker.
Musca praecox, Walker.
Musca angustifrons, Thomson.
Musca bivittata, Thomson.
Musca mveisquama, Thomson.
Musca eutaeniata, Bigot.
Musca promisca, Awati,

This strikingly marked species is the most important tropical house, bazaar
and camp fly. It is of peculiar interest, as in its habits it links the house-fly with
the wild haematophagous species. Its larvae may be found in a great variety of
food-stuffs, in isolated patches of cow-dung, horse-dung and human excrement ;
also in cow-dung and horse-dung when piled in heaps, in “night- soil trenches, and in
decaying vegetable matter. I have also bred it in dog’s dung and in the decomposing
bodies of rabbits. As it commonly breeds in patches of cow-dung when dropped
in the field, the adults are frequently found on cattle and horses far from human
dwellings. Under these conditions they feed on the discharge coming from sores,
and from the eyes and noses of cattle. As often as not, the specimens which hatch
out of larvae in isolated patches of cow-dung are very small, and they are most
irritating pests in the fields, swarming about one’s head, as do the adults of Hvdrotaea
ivvitans in the summer months in Europe.

In India it swarms in the bazaars, where it may be seen on food-stuffs of all kinds,
as well as on human beings, particularly around the eyes of children. In Mesopo-
tamia it was quite common to see the eyes of small children covered with the females
of this species, and this explains how the eyes of adults become infected with various
pathogenic bacteria,

418 MAJOR W. S. PATTON.

Musca humilis is very widely distributed, probably more so than any other
species of the genus. I have seen specimens from many parts of Africa, Palestine,
Mesopotamia, Persia, and from the whole of India to China in the East.

The late Professor Stein examined Wiedemann’s type of humilis, as well as
Thomson’s types of angusiifrons and bivittata, and says all three are the same species.
Until I can for myself examine these types I accept this determination.

I have examined Walker’s type of Musca conducens, a male collected by Wallace
at Macassar, Celebes ; it is in good condition and is a small example of M. humilis.
Walker’s type of AM. praecox, a headless male from Ceram, Malasia, is exactly similar,
and is also a small specimen of M/. humilis. Walker's type of IZ. primitiva, a female
from Fu-chau-fu, South China, is a typical specimen of M. humilis.

Bigot’s collection of Musca, fifty-three specimens, contains six males of his Musca
eutaeniata from “ the Indies,” chiefly from Pondichery and Cochin China, all of
which are rather small, but typical specimens of M. humilis. It will be remembered
that Brauer examined these specimens, and came to the conclusion that like eight
of the other specimens of Musca in Bigot’s collection (Brauer’s numbers 257 to 265,
inclusive) they were ‘“ Musca ead.n. (wie die ersten 9 wohl domestica). ~ sit is quite
clear from this that Brauer had a very vague knowledge of the species of the genus
Musca, otherwise he would never have called these six specimens of M. humilis
(eutaeniata), domestica. This is a warning not to accept the determination of types
by others as final without oneself examining them. Stein states that M. mveisquama,
Thomson, is identical with J/. domestica, L. Major Austen has, however, drawn
my attention to Thomson’s description of this species, in which he clearly states
that mivetsquama is similar to his angustifrons, and has two black thoracic stripes.
It is quite possible that miversquama is Musca pumila, Macquart.

Awati gives angustifrons as a synonym for his fusca promisca, and his description
of this species makes it quite clear that it is MW. humilis. This author, when describing
the three species, Musca nebulo (multispina), M. domestica atypical (divaricata), and
M. humilis (promisca), states that in the case of mebulo there are “‘ two or more spines
on the humeral vein which may be present on both wings or on one of them only ; ’
in the case of MW. domestica atypical (divaricata), he says there is “‘ only one spine on
the humeral vein,’ and in M. humilis also only one spine on this vein. In his
remarks at the end of the description of M. divaricata he says: ‘‘ There are always
more than one spine on the humeral vein in Musca multispina, whereas in Musca
divaricata there are never more than one.’ By the humeral vein Awati evidently
refers to the large basal root vein from which arise the subcostal and radial (subcostal ;
Ist longitudinal) veins just before the smal] humeral transverse vein joins this root
vein to the costal. In all the species of Musca which I have examined, there
are always one or more small curved bristles situated on the inner side of the vein.
I have made a careful examination of the number of bristles present on the veins of
hundreds of specimens of Musca nebulo, M. domestica (atypical) and M. humilis, and
in a long series of domestica (atypical) in the National Collection a large percentage
had two bristles.on the right root vein, and only one on the left ; a few, on the other
hand, had two on the left vein and only one onthe right. The majority, however,
had only one bristle on each vein. In the case of a long series of nebulo, I find that
a few have two bristles on the left vein and one on the right, while some have two
on the right and only one on the left. But, and this is important in view of Awati’s
statement that in this species there are always two bristles on this root vein on both
sides, I find that the majority of the specimens of mebulo only have one bristle on
each root vein; this also applies to humilis.

Musca humilis can always be recognised by noting that it is a much greyer fly
than the two others noted above ; that in the female “the black thoracic stripes are
united behind the suture, forming one broad stripe, but are separated in front of
the suture ; in the male the stripes on each side have coalesced to form one broad

SPECIES OF THE GENUS MUSCA, LINNAEUS. 419

black stripe, which diverges posteriorly away from the central grey stripe; the
abdomen in the female is quite unlike that of the male, and is yellowish grey with
black stripes and bands; that of the male yellowish with silvery patches and a
central black stripe. The frons of the female is wide, that of the male a little narrower
than the frons of the male nebulo.

It would be interesting to know if this species has found its way to any part of
the New World.

4. Musea sp. incerta (‘‘ Textbook of Medical Entomology,’ Patton and Cragg,
p. 334).

This distinct species was bred in 1910 from larvae collected from the contents
of the stomach and intestines of sheep slaughtered at the slaughter-house, Saidapet,
Madras, and since then another good series has been obtained from the same source ;
it also breeds in horse-dung. It will be described as a new species in a forthcoming
paper on some new Indian species of the genus Musca.

On a superficial examination it may be mistaken for Musca nebulo, but it will
be noted that the ground-colour of the thorax is darker and is of a bluish colour.
It has four black thoracic stripes, and the male frons is much narrower than that of
the male webulo, the eyes being separated only by a fine black line. The abdomen
in both sexes is much more orange-coloured, and in the male there are no marginal
silvery patches on the apparent second segment ; in the male nebulo these silvery
patches are always well marked.

Musca pumila, Macquart.
Synonyms: Ausca minor, Macquart.
Musca vetustissima, Walker. ‘
Musca humilis, Stein (mec Wiedemann).
Musca corvina, Froggatt (nec Fabricius).

I first bred this species from larvae collected from human excrement in 1910
in Madras, and in 1920 obtained a large number from the same source. Mr. Senior-
White has collected it in Ceylon. I have received a number of specimens from
Australia, sent me by Mr. Tay lor, Mr. Froggatt, Mr. Hill and Dr. Ferguson, and
all are identical with the species in the National Collection determined by Major
Austen as Musca pumila. There is also a specimen in Bigot’s collection from
Australia labelled Musca pumila, Macquart. As it is superficially like MW. humilis
it has been mistaken for that species. In both sexes the thoracic markings are exactly
similar to those of WZ. humulis, but the ground-colour of the thorax is markedly blue,
whereas in humilis it is yellowish grey. The abdomen in the female is also bluish,
with black stripes and bands. a far I have not seen the typical Musca humilis
from Australia, and Musca pumila seems to take its place.

6. Musca ventrosa, Wiedemann.
Synonyms: Musca xanthomela, Walker.
Musca pungoana, Warsch.
Musca nigrithorax, Stein.
Musca kasauliensis, Awati.
Musca hill1, Johnson & Bancroft.

This small species, with its dark thorax with four stripes and orange-yellow
abdomen, is a true haematophagous Musca, and is mainly seen on animals, and on
foliage near them. Wiedemann’s types of ventrosa came from Sumatra and China.
I have examined the type, a female, of Walker’s Musca xanthomela from Macassar,
Celebes, and find it is a typical specimen of ventrosa; Stein’s nigrithorax from
Samarang and Batavia, and Karsch’s Musca pungoana from Pungo Ndongo,
Portuguese West Africa, are also this species.

420 MAJOR W. S. PATTON.

I have examined the paratypes of Musca hilli, Johnson & Bancroft, and except
for some dark markings on the apparent second and third abdominal segments, they
are identical with the Indian specimens of ventrosa, in many of which there are also
some dark markings on the same segments. I can see no reason at present for
considering M. hilli to be a distinct species, but hope to compare the external
genitalia of the Australian and Indian specimens to settle its identity.

Awati gives his Musca kasauliensis as a synonym of M. nigrithorax, and there
seems to be no doubt that his species is M. ventrosa.

Musca ventrosa is widely distributed in India, Burma, Assam and Ceylon, and is
also common in Africa; it breeds in cow-dung.

7. Musca albomaculata, Macquart.

Synonyms : Musca dorsomaculata, Macquart.
Musca convexifrons, auctores (nec Thomson).
Musca setigera, Awati.

This species was determined for me by Professor Bezzi as Musca convexifrons,
Thomson. It is one of the common haematophagous species, and has been fully
described in collaboration with Major Cragg, I.M.S. On comparing Thomson's
description of convexifrons with it I have come to the conclusion that the Indian
species long known under this name is not Thomson’s species. The discrepancies are
as follows: Thomson says that his specimen, a male from China, measures 6 mm.,
but the largest Indian specimens I have seen measure only 5:5 mm. ; and further,
he says that his species is like Musca autumnalis, de Geer (corvina, F.), which is quite
unlike the.Indian species. In order to settle this point I recently sent a typical
male to Professor Sjéstedt for comparison with Thomson’s type, and Dr. Roman
has kindly given me a note as the result of his examination and comparison of the
two species ; from this it is quite clear that the Indian species is not Musca convext-
frons. Dr. Villeneuve tells me that he has compared this Indian Musca with
Macquart’s types of Musca albomaculata and M. dorsomaculata and finds they ‘are
identical. I accept this determination for the present. I have very little doubt
that Thomson’s convexifrons is the species known as Musca fergusont, Johnson &
Bancroft (australis, Macquart, nec Boisduval), a species which is widely distributed
in Australia, as well as in the many neighbouring islands, and presumably extends
to China. At any rate, this species is very like Musca autumnalis, de Geer. If this
should eventually prove to be the case, the Australian species must in future be
known as Musca convexifrons, Thomson.

8. Musca pattoni, Austen.

Synonym: Musca spinosa, Awati.

This species has been fully described by Austen, and the types are in the National
Collection. It is widely distributed in India, Burma, Assam and Ceylon, but I have
not seen any specimens from any locality outside India. Awati has described the
female under the name of Musca spinosa.

9. Musca gibsoni, Patton & Cragg.
Synonym: Musca latiparafrons, Awati.

This species was described by me in collaboration with Major Cragg, I.M.S. It
is especially abundant in hill stations on animals. The female has been described
by Awati under the name Musca lattparafrons.

SPECIES OF THE GENUS MUSCA, LINNAEUS. 421

10. Musca spinohumera, Awati.

The female of this larviparous species has been fully described by Awati.
Mr. Senior-White recently collected a large number on cattle at Pusa and noted
that it is haematophagous in habit. He also observetl that it deposited one larva
at a time, thus confirming Awati’s observations ; the puparium is very like that of
Musca pattoni, and is of a dirty white colour.

I have examined this collection, which also contains the male, and compared
both sexes with those of MW. pattont and with the types of M. mesopotamiensis, Patton.
Though M. spinohumera superficially resembles M. pattoni, they are quite distinct,
the one being oviparous and the other larviparous. Musca spinohumera is, however,
closely related to W. mesopotamiensis, but there are some good characters by which
they can be distinguished, and here again one is oviparous and the other larviparous.
I have not seen M. spinohumera from South India, and it appears to be peculiar
to Northern India.

Awati mentions /. corvina, Fabricius, as a synonym of his spinohuwmera, evidently
meaning the form which the late Professor Portchinsky referred to as Musca corvina,
Fabricius (vivtpara). It will be remembered that Portchinsky pointed out that there
were two types of M. corvina in Russia, one which he called MW. corvina ovipara,
found in Northern Russia, which lays stalked eggs and has a white puparium, and
the other, WV. corvina vivipara, which deposits larvae and has a brown puparium,
and is only found in the Crimea. In the paper on “ Certain haematophagous species
of the Genus Musca” (Ind. Jl. Med. Res., i, no. 1, 1913), in collaboration with Major
Cragg, I.M.S., attention was drawn to the observations of Portchinsky in a footnote
on page 12, and after examining a good series of both forms, it was concluded that
they were distinct species. To quote our words, “‘ From our knowledge of the
Oviparous convexifrons (spined eggs and white puparium), patton (spined eggs and
dirty grey puparium), and the larviparous bezzi1, it appears to us that Portchinsky
has confused two distinct species of Musca, one entirely oviparous and the other
entirely larviparous.’’ Since then I have studied all the stages of the oviparous
species now known as Musca autumnalis, de Geer, in the South of England, and
observed that it lays its stalked eggs singly in patches of fresh cow-dung dropped
in the field, and that the puparium is white. But with regard to the larviparous
species, Major Austen has drawn my attention to the curious fact that there is no
description of it in any of Portchinsky’s writings, and yet I find that Schnabl and
Dziedzicki in.their work, ‘“‘ Die Anthomyiden,” speak of it as Wusca lavvipara,
Portchinsky (olim Musca corvinoides, in litt. Schnabl & Dziedzicki). As they
partly describe the male, the species must in future be known as Musca larvipara,
Schnabl & Dziedzicki, and not Portchinsky.

As my material of this species is old and not in good preservation, I shall be
glad to exchange any Indian species for fresh material.

11. Musea hezzii, Patton & Cragg.

Synonym: Musca pilosa, Awati.

This large handsome fly is very common in all the Indian hill stations, and also
at the foot of the hills. It is purely haematophagous in habit, and is only found on
and around animals. The female deposits one larva at a time, in the second stage,
on patches of fresh cow-dung.

M. bezzii belongs to the lusoria group, but is distinct from that species. The
male has a dark thorax and yellowish abdomen, with silvery patches and stripes.
The female is greyish, with black bands and stripes.

422 MAJOR W. S. PATTON.

. Musca (Pristirhynchomyia ; Philaematomyia) lineata, Brunetti.

This small haematophagous species is widely distributed in India, and is always
found on and near cattle in the fields. It is not a true blood-sucker in the sense that
it can draw blood, but it has moderately well-developed prestomal teeth, and can
certainly scratch a clot off the skin and suck up any fluid under it. It thus shows
an early change towards the true scratching proboscis, so well seen in the case of
Musca crassirostris, Stein, which is able to draw blood with the aid of its prestomal
teeth.

It will be remembered that in the typical /ineata, the rather narrow thoracic
stripes are distinctly separated, whereas in what appears to be a variety which |
bred in Coonoor from cow-dung, and which Mr. Senior-White has also bred from the
same source in Ceylon, the thoracic stripes are distinctly broader, and in the male
almost, 1f not entirely, coalescent behind the suture; in addition, the front of the
males of this v ariety is much narrower than in the ty pical lineata. \am not able to
express a definite and final opinion as to the identity of this variety, but as I have
the larvae of both the species, as well as a large number of adults, I hope to do so
shortly. I may, however, point out that the male is quite easily mistaken for a small
specimen of Musca humilis, but it is certainly not that species.

13. Musca cingalaisina, Bigot.

Synonyms: Musca pollinosa, Stein.
Musca (Philaematomyia) indica, Awati.

I have examined the type of Bigot’s cingalaisina, a female without a head, from
Ceylon, and have no hesitation in saying it is a typical specimen of Musca indica,
Awati. I have examined hundreds of specimens of this most interesting species
from many parts of South India and from Ceylon collected by Mr. Senior-White.
It is extremely common in Ceylon, and though neither Mr. Senior-White nor I have
ever caught it on cattle in the act of sucking blood, it-is unquestionably a blood-sucker,
and can draw blood with the aid of its prestomal teeth. Mr. Senior-White is now
engaged in studying its feeding habits, which appear to be very peculiar. In localities
where it occurs it can be caught i in large numbers sitting about on leaves, etc., and
on fresh cow-dung, on which it deposits its third-stage larva. Itis very curious that

although it can be seen on cow-dung close to a cow, it has never been seen either by
ie ecniee -White or myself actually on the animal. Awati also states he has not
seen it sucking blood. ‘

I have no doubt whatever that Stein’s follinosa is this species, andit appears to
be common in Batavia, Samarang and Tandjong Priok, in the Dutch East Indies.
Brauer came to the conclusion that the type of cimgalaisina was a specimen of Musca
domestica, a remarkable determination. -

14. Musca (Ptilolepis) inferior, Stein.
Synonym : Philaematomyia gurneyi, Patton & Cragg.

In a recent paper Bezzi has made this species the type of a new genus, Ptilolepis,
mainly basing it on the presence of dark hairs on the inner portions of the squamae.
He also gives some additional characters, the majority of which, however, are in my
opinion not generic characters, but are common to many of the other species of Musca.
He mentions “Eyes bare, rather distant in the male. Parafrontal hairs of the
female arranged in more than one row,” and other characters based on chaetotaxy.
The presence, absence and arrangement of bristles and hairs are not reliable characters
on which to base genera. Further, I would point out that the characters of the
proboscis, such as, “ thickened basally with chitinous terminal teeth,’”’ are common

SPECIES OF THE GENUS MUSCA, LINNAEUS. 423

to several species, and that there are three bristles on the radial root vein and not
one, as mentioned by Bezzi. It is true, as Bezzi points out, that the presence of long
hairs on the posterior portion of the upper surface of the squamae is unique in this
species, but I do not consider this to be of generic importance.

Musca inferior is a true blood-sucker and is widely distributed in India, but it
is never seen in large numbers and is easily missed. It can always be recognised
by noting that as soon as it settles on the skin of an animal it begins to suck blood
at once, whereas Musca bezzit, which closely resembles it, flits about from one spot
to another. I have one male specimen, which was caught on a patch of human
excrement. It probably breeds in cow-dung and is almost certainly larviparous.

15. Musca (Philaematomyia) crassirostris, Stein.
Synonym : Philaematomyia insignis, Austen.

Musca crassirostris is one of the most important cattle pests in India, and it would
be well to name it the “Cattle Fly.” I have little doubt that it very materially
reduces the quantity of milk of Indian milch cows, owing to its vicious biting habits
and incredible numbers. I have repeatedly seen the legs of a cow covered with
large numbers of this fly, and the animal then spends a great amount of its time
driving them off. These animals are often forced to lie down and curl their legs
under their bodies, in order to protect themselves from the attacks of IW. crassirestris.

It is interesting to note that it lays a relatively large egg, which hatches out in
a very short time, suggesting that some development has already commenced while
the egg was still in the ovary.

It is widely distributed in the Oriental Region, and is also found in many parts
of Africa, and I have little doubt that when it is more carefully looked for will be
found in many other localities. I see no reason for placing this species in a distinct
genus, Philaematomyia, on the structure of the proboscis alone, for I now know
five species that have this type of proboscis, which are otherwise typical specimens
of Musca.

In a paper that is now in preparation I hope to describe five new species of Musca
from India (inclusive of No. 4 noted above), one of which has well-developed prestomal
teeth. I have now received specimens of the typical Musca domestica from Kashmir,
as well as M. tempestiva, Fallén, and M. vitripennis, Meigen, all collected by Dr. Baini
Prashad ; and including these, there are now 22 good species of the genus Musca
found in India. So far, I have not seen a single specimen of M. albina, Wiedemann,
from any part of India, although the type is said to have come from the East Indies.
This species has been sent to me from Egypt, and Bezzi records it under the name
speculifera from Djerba, Tunis. I have not seen any species which could be identified
as Musca minuta, Awati, M. negriabdomina, Awati, or M. siriatecta, Awati. I
shall be very glad of any specimens of Musca from any locality in the Oriental Region,
and hope that those who have opportunities of collecting these flies will do so and
send them to me, so that the revision of this group may be satisfactorily completed.

THE AUSTRALASIAN SPECIES.

Although the National Collection is not very rich in material from the Australasian
Region, it contains several types and paratypes, which have enabled me to settle
the identity of some of the doubtful species. I have also been able to supplement this
collection with a number of specimens of most of the species sent me from time to
time by Mr. F. H. Taylor, Mr. W. W. Froggatt (Entomologist to the Government of
New South Wales), Dr. Eustace W. Ferguson (Department of Public Health, Sydney),
Professor T. Harvey Johnson, and Mr. G. F. Hill (Entomologist to the Australian

424 MAJOR W. S,. PATTON.

Institute of Tropical Medicine). Mr. Hill in particular recently sent me a valuable
collection of MuscipAE from various localities. I wish to take this opportunity
of thanking these gentlemen for the trouble they have taken in collecting and sending
all these specimens.

The revision of the species from the Australasian Region has been most difficult,
owing to the great confusion existing as to the correct names of even the commonest
species ; most, if not all, of these have been described by the older authors, but
unfortunately their descriptions are for the most part valueless, if not actually mis-
leading. It is for this reason that I would earnestly appeal to those who have oppor-
tunities of collecting species of Musca, to let me have as many specimens as possible.
I would particularly like large numbers of house-flies from any part of the Region,
either in 80 per cent. alcohol or packed in matchboxes. It will only be by comparing
large numbers of specimens of the species from as many localities as possible with
those from other regions, particularly the Oriental, that it will be possible to settle
the true identity of the species of the older authors. And it would be most unfor-
tunate if my final revision were to be incomplete merely for the want of sufficient
material. Larvae and flies bred from them are the best material for the comparative
studies I now have in hand.

1. Musca domestica, L. (typical).
Synonym: Musca vicaria, Walker.

Of the 58 specimens of this species in the National Collection from various parts
of Australia, and from the Sandwich, Samoan, Solomon and Fiji Islands, I have
provisionally determined 21 (including the type of Walker’s vicaria, a female from
New Zealand), as belonging to this species. And I have specimens in my own
collection which are undoubtedly this species.

2. Musca domestica, L. (atypical).
Synonym: Musca antiquissima, Walker.

The remaining 36 specimens in the National Collection (including the type of
Walker’s Musca antiquissima, a male), belong to the form of domestica in which the
male has a front much narrower than that of the typical form. Most of the
specimens in my own collection conform to this type, so that it would appear that
it is one of the common house-flies of Australia; its true identity will only be
arrived at by examining microscopic preparations of the external genitalia of both
sexes and comparing them with the similar form from other regions. I should be
glad of some hundreds of specimens of this species.

3. Musca pumila, Macquart.

Synonyms: Musca minor, Macquart.
Musca vetustissima, Walker.
Musca autumnalis (corvina), Froggatt (mec de Geer).
Musca humilis, Stein, Bezzi (nec Wiedemann).

This species is identical with a Musca bred by me more than 10 years ago in
Madras. It has been confused with Musca humilis, Wiedemann. It appears to be
a troublesome bush fly in Australia, and I have four small females sent me by
Mr. Froggatt with a note that they were caught in tents. There are three specimens,
one male and two females, in the National Collection from Cloncurry, Queensland,
collected by Dr. Priestly, who notes that it is a troublesome fly, settling on the
human eye and probably carrying the bacteria of eye diseases. In Australia it
appears to have taken the place of Musca humilis. The Australian specimens differ
slightly from the Indian in that the frontal stripe of the male is usually a little
broader, but this is a variable character.

SPECIES OF THE GENUS MUSCA, LINNAEUS. 425

4. Musca terrae-reginae, Johnson & Bancroft.

I have seen only six specimens, five females and one male, of this species, sent
me by Mr. Hill and Prof. Johnson, as well as the paratypes deposited in the
National Collection by the latter. I am not able with this very meagre material to
come to a definite conclusion regarding the true identity of this species. It will be
impossible to discover what Musca prisca, Walker, really is, as the type (a female
from New Zealand) is unfortunately a greasy specimen, and was evidently in this

condition when Walker described it; it should never have been described. One
thing is certain, it is aspecies with four black thoracic stripes and may quite well be
Musca terrae-reginae. I hope to re-examine it more critically, and to compare
it with M. terrae-reginae, but if this examination does not lead to any definite
conclusion, I propose to drop this name altogether. More specimens of M. terrae-
veginae ate, however, required in order to compare it with some of the Indian species
with which it is allied.

5. Musca ventrosa, Wiedemann.

Synonyms: Musca xanthomela, Walker.
Musca pungoana, Warsch.
Musca nigrithorax, Stein.
Musca kasauliensis, Awati.
Musca hilli, Johnson & Bancroft.

In my notes on the Oriental species I pointed out that I had examined the para-
types of Musca hilli, Johnson & Bancroft, deposited in the National Collection by
Prof. Johnson, and considered it to be identical with Musca ventrosa, Wiedemann.
Mr. Hill recently sent me a long series of M. hill1, and among them a couple which
have been determined by Prof. Johnson, and though most of these specimens show
varying amounts of dark stripes on the apparent third and fourth abdominal segments,
they appear to be identical with Indian specimens of MW. ventrosa. The final deter-
mination will depend on a comparative study of the external genitalia of both sexes,
which I now have in hand. It may be that Musca hilli is a good species.

6. Musea convexifrons, Thomson.

Synonyms: Musca australis, Macquart (nec Boisduval).
Musca fergusont, Johnson & Bancroft.
Musca lusoria, Bezzi (nec Wiedemann).

In my notes on the Oriental species I have pointed out that Musca fergusoni,
Johnson & Bancroft, is the species known as Musca convexifrons, Thomson. ‘It
will be remembered that Thomson described as his type a male from China, and that
for a long time it was believed to be identical with a common Oriental species, which
is recorded above under the name of Musca albomaculata, Macquart. But I was
convinced that the Oriental species was distinct from Thomson’s M. convexifrons,
and was for a long time puzzled as to the identity of Thomson’s species. I now,
however, have no doubt whatever that it is the common Australian haematophagous
Musca recently re-described by Johnson and Bancroft under the name Musca
Jergusont. Mr. Hill recently sent me a long series, some of which were collected on
Palm and Magnetic Islands, and I have been able to compare it with Musca lusoria,
Wiedemann, and Musca bezzii, Patton & Cragg. In Musca convexifrons there are
normally two bristles on the basal portion of the radial vein, sometimes three ; in
M. lusoria there are normally four bristles, sometimes more and sometimes less ;
andin M. bezzii there are normally five. M. convexifrons differs in many other respects
from both these species.

426 MAJOR W. S. PATTON.

These six species of Musca represent all those known to me from the Australasian
Region, but I have no doubt that there are several others which have yet to be
discovered. It is to be regretted that I have no specimens of Musca from any part
of New Zealand, and I trust that those who are interested in this important group
of flies will send me material from this part of the region.

In my next series of notes I shall record my studies of the species of Musca from
the Ethiopian Region, a very large collection of which Dr. Guy A. Kx. Marshall has
kindly placed at my disposal.

427

MOSQUITO LIFE IN SURREY DURING 1921.
By Lt.-Col. S. P. James, I.M.S. (retd.),
Ministry of Health.

Mr. Malcolm E. MacGregor’s interesting paper, ‘‘ The Influence of Drought
upon Mosquito Life in Surrey ”’ (Bull. Ent. Res. xii, p. 205), has led me to examine
our records of observations made in that county in connection with inquiries into
indigenous malaria in England. As regards certain areas of the county during the
past year, our experience of the scarcity of some kinds of mosquitos is the same
as Mr. MacGregor’s, but (if it is desired to draw conclusions for the county as a
whole) it seems important also to note that there are other areas in which the findings
differ from those in the particular locality upon which he reported. Dorking, Epsom
and Arbrook Common, which are within a few miles of Wisley, are examples of
such areas. In these and some other localities in the county we have obtained
larvae of A. bifurcatus without difficulty throughout the year. At Epsom, large
numbers of larvae of this species in the fourth instar were found early in February
this year in a well-shaded permanent pool, which we have examined regularly since
1917, and adults were caught in the open in April. Larvae were numerous
throughout the year, and adults, both male and female, were captured in the open
as late as 22nd October. At Dorking, larvae of bifwrcatus were plentiful in several
natural collections of water throughout the year. One of them—the “ Mill Ponds
stream ’’—is only 100 yards from a row of houses ; another—the ‘‘ Stonebridge
stream '’—yielded bifurcatus larvae on 19th August at every place examined along
a stretch of 400 yards. On the same day many larvae of this species were found,
along with larvae of C. pipiens, in the rainwater barrel of a house about 200 yards
from the stream.

Next, as regards A. plumbeus and Finlaya geniculaia, we have collected larvae
of both species from tree-holes at Dorking and Box Hiil on many occasions since
the middle of August. Again, the incidence of Theobaldia annulata and Ochlerotatus
nemorosus at Dorking and other localities differed this year from that in
Mr. MacGregor’s area. Adults of Theobaldia annulata were caught in a bedroom at
Dorking during July, and larvae have been plentiful in a large stagnant ditch
throughout the year. Also it is worthy of note that my laboratory assistant
(P. G. Shute) collected from Arbrook Common in June a larva of a species (Aédes
cinereus) which is not included in Mr. MacGregor’s list of the species found at Wisley.

As regards A. maculipennis, Mr. MacGregor records the important observation
that, when nearly all the available breeding-places in his area were dry, there was
an increase in the actual numbers of larvae found in the breeding-places which
remained. As well as similar findings, our records contain the observation that this
year the river Wey, at Guildford, for the first time since we began to examine it in
1918, harboured many maculipennis larvae. During July and August, my laboratory
assistant, using a boat, collected numerous specimens along both banks of the river
over a distance of a mile.

This and other observations in Surrey and Kent lead us to believe that the
presence in our area of permanent breeding-places of various kinds sufficiently explains
the differences between the records for our area and for the area examined by
Mr. MacGregor. In India, where long periods of drought recur annually, the relative

(5298) 21

428 LT.-COL. S. P. JAMES.

importance of permanent and temporary breeding-places has been worked out in
detail. The account on pages 82-83 of Major Christophers’ report on “‘ Malaria in
the Punjab’ shows clearly, I think, that what Mr. MacGregor has described as
happening in his area this year is what happens each year in certain large tracts of
that country. With respect to the various mosquitos which are indigenous in an
area, Major Christophers pointed out that, however hard pressed by drought some
of them may be in a particular part of the area, it is always possible to find, within
their usual distance of spread, some permanent collections of water in which they
are breeding freely. Also, that when temporary breeding-places reappear, the
various species spread widely from these permanent sources (which he calls
“mosquito sanctuaries ’’) until in a short time they can be found again throughout
the area. This explains the well-established Indian observation that a particularly
dry year has no permanent effect in reducing Anopheles mosquitos. In the
Punjab the chief Anopheles “ sanctuaries ’’ were found to be :—(1) Large river-beds
like those of the Jumna, Beas, etc.; (2) jheels and large tanks filled with aquatic
vegetation, with their associated swamps and pools; (3) extensive brickfields and
large excavated pits near towns and villages ; (4) irrigation systems.

Nearly the same headings would be applicable to the permanent mosquito
‘sanctuaries ’’’ in Surrey and Kent. As some of them exist within a few miles of
the area reported on by Mr. MacGregor, a probable answer, based on Indian
experience, can already be given to his question whether the species dealt with will
be rare in his locality during future years.

* Scientific Memoirs by Medical Officers of the Government of India, New Series No. 46, 1911.

429

A NEW APHID GENUS AND SPECIES FOUND IN ENGLAND,

By FRED. V. THEOBALD.

Genus Laingia, nov.

This marked genus appears to be intermediate between Atheroides, Haliday,
and Sipha, Passerini.

The characters are as follows: Body elongate in the apterous female. Head
somewhat rounded in front to almost flat, moderately large ; no frontal or antennal
tubercles. Eyes large; ocular process prominent (fig. 1, E, oc.p.) and truncate.
Proboscis (C) rather short and thick; last two segments of about equal length.
Antennae (A) rather short, of five segments, reaching to or just past the pronotum (B).
Thoracic segments large and distinct. Legs rather short and thick, the prothoracic

Fig. 1. Laingiapsammae,sp.n.: A,antenna of apterous viviparous 9 ; B, head
and prothorax; C, proboscis; D, anal plate (a.p.), cauda (c), apicalsegment (8) ;
E, eye, ocular process (0c.p.); F, cornicle.

pair far from the mesothoracic. Cornicles (I) round, very slightly raised.
Cauda (D, c) rather small, broad, parallel-sided and slightly convex apically. Anal
plate (D, a.p.) somewhat rounded apically, broader than cauda, sides straight but
divergent ; both anal plate and cauda hidden by the prominent semicircular apical
segment of the body (D, ’). Integument finely spinulose. A few short hairs on
the body ; long, thicker ones on head and apex.

This genus differs from Szphain the non-knobbed cauda and from Atheroides in
the non-rounded cauda. Superficially it bears some resemblance to Thripsaphis,
but the ocular processes at once separate it from that genus.

I have named it after Mr. F. Laing, of the British Museum, who pointed out to
me that it appeared to be intermediate between Atheroides and Sipha.

Laingia psammae, sp. nov.

A pterous viviparous female.—Elongate, rather narrow. Colour varying from dull
straw to dingy brownish-green. Eyes dark. Legs short, rather thick, darker than
body. Abdomen with dusky lateral patches and small dusky spots ; apical segment
dusky, also apex of cauda ; a dusky area around base of cornicles ; anal plate dusky.

(5296) 212

430 FRED. V. THEOBALD.

Antennae of five segments, as long as or a little longer than head and pronotum ;
Ist segment wider and longer than 2nd ; 3rd as long as to a little longer than 5th ;
4th small, from one-fourth to a little more than the length of the 3rd ; 5th with basal
area as long as to alittle longer than the 4th; four curved hairs on basal segment ;.
two on 2nd; three to four on 3rd; one on 4th near apex, opposite the sen-
sorium ; two on apex of basal area of the 5th; one primary and several secondary
sensoria at junction of basal area and flagellum ; segments 3-5 imbricated, the last
markedly so; at apex three small hairs. The more or less rounded head bears six
long thick hairs in front and several shorter ones passing on to the vertex. Eyes |
large, with prominent quadrate ocular processes. Proboscis rather short and thick, —
reaching to the second coxae; apical and penultimate segments about equal in
length, both dusky. Integument finely spinulose ; a few short hairs on body, longest
towards the apex. Apical segment rounded, with four long hairs and some short
ones, completely covering the cauda. Cornicles round, on slightly elevated cones,
which are surrounded by a small dark area. Cauda short and broad, convex at
apex, finely spinose, with two long apical hairs ; dark, especially apically. Anal plate
broader than cauda, sides straight but divergent. Legs rather short and thick,
a few hairs on femora ; more on tibiae ; pro- and mesothoracic pairs widely separate.

Length, 1-9-2-2 mm.
Locality. Littlestone, Kent (vii.1921).

Food-plants—Marram grass (Psamma arenaria) and meadow foxtail grass. |
(Alopecurus pratensis). |

Found first by Captain A. Duffield, M.C., in great abundance on the marram
grass growing on the sandhills near Littlestone. It lives in the blossom heads, |
usually deeply hidden in the heads, being of much the same colour as the ripening
blossoms and consequently not easily detected unless in large numbers. Whilst
visiting Littlestone in September I could find no trace of living aphides, but all the
ripe grass was smothered with their exuviae and black soot fungus. Mounted exuviae
from meadow foxtail grass on the marsh near by show that it had also served as a
food-plant. This Aphid produces much honey-dew, which attracted various insects.
It was preyed upon by countless ladybird beetles, the chief being Adalia bipunctata,
and also by many small Syrphid larvae. These, however, had not prevented great
numbers of alatae from appearing, for I found large quantities of the cast nymph skins.
The grass seed seemed to have been quite ruined by the swarms of this plant-louse.

431

SOME SIAMESE TABANIDAE.
By Major E. E. Austen, D.S.O.

The potentialities of Siam as a field for discovery have hitherto been almost
entirely neglected by collectors of Diptera, and in consequence, apart from an
occasional description of a new species, little or nothing has been published upon
the Siamese representatives of this Order. It follows that the only available method
of making determinations of such material is the exceedingly laborious and
wearisome. one of reading through descriptions of the older authors like Wiedemann
and Macquart, which, though based on types collected in other parts of the Oriental
Region, may possibly be found to apply to one or other specimen from Siam.

The following little paper has been prepared in the hope not only of doing something
to facilitate the study of Siamese TABANIDAE, but also of arousing interest among
those who have the opportunity of collecting these insects within the confines of the
kingdom of our eastern ally, so that our knowledge may ere long be extended. The
fact that the present paper records only a single species in the case of Chrysops,
and two species in that of Haematopota, is sufficient indication of its incompleteness ;
of the latter genus, at any rate, it is almost certain that many more than two
species are to be found in Siam.

The basis of the paper is a small but interesting collection of Siamese TABANIDAE,
recently formed and presented to the British Museum (Natural History), by Dr. M. E.
Barnes, of the Rockefeller Foundation, New York, and now of Bangkok. In expressing
to Dr. Barnes the grateful thanks of the Museum for his kindness, the author desires
to draw attention to the interesting field-notes accompanying many of the specimens,
which greatly enhance the value of the collection. As a stimulus to others towards
further effort in the same direction, it may be pointed out that, of the seven species
of Tabanus obtained by Dr. Barnes, no fewer than five appear to be new.

In order, so far as possible, to extend the scope of the paper, the small amount of
Siamese Tabanid material previously in the National Collection has been studied
in conjunction with Dr. Barnes’s specimens and recorded, or, where necessary,
described, in the following pages. Unless otherwise stated, however, every specimen
mentioned was collected by Dr. Barnes.

It is only necessary to add that the types and paratypes of all new species included
in the present paper are in the British Museum (Natural History).

PANGONIINAE.

Genus Chrysops, Mg.

Chrysops flavecincta, Ric.

Chrysops flavocinctus, Ricardo, Ann. Mag. Nat. Hist. (7) ix, p. 380 (May 1902).

Three 99, Doi Chom Chang, near Chiengmai, 16.iv.1921. With reference to
these specimens, Dr. Barnes writes: ‘‘ Caught on Doi Chom Chang, while attacking
me ina cottage. These flies are very common on the mountain in that region, and
are extremely troublesome at a sanatorium situated at an altitude of about 2,500 ft.
They were never reported from the top of the mountain (5,500 ft.) until a few cottages
were built there, but since that time they are to be found on the peak also.

432 MAJOR E. E. AUSTEN.

“ They attack very cautiously, approaching their victim from behind, and rarely
making for the face. In flight, they somewhat resemble ‘ Hover-flies,’ noiselessly
approaching and settling upon the ears, back of the neck, legs, back of the hands,
and on the exposed elbows. It is only the severe pain of the bite that attracts the
victim’s attention, and should the sufferer make the slightest movement, the flies
at once effect their escape.

“The wound made by their bite is very painful, and causes much swelling in
many people. A minute bleb forms at the site of the puncture, and if the contents
be expressed the irritation usually subsides, though the pain and intense itching may
persist for several days. If these wounds become infected, as is frequently the
case in children, who are very apt to scratch them, they heal very slowly.

“This insect is by far the most troublesome biting fly in the Doi Chom Chang
RESIONs

Specimens of Chrysops flavocincta already in the National Collection show that
the area of distribution of this species, which is one of the smallest of the Oriental
representatives of its genus at present known, includes Ceylon, North-eastern India
(Khasi Hills, Assam), and Borneo (Sarawak).

TABANINAE.

Genus Haematopota, Me.
Haematopota pachycera, Big.
Haematopota pachycera, Bigot, Nouv. Archiv. Mus. Hist. Nat., Paris (3) ii,
p. 206 (1890).
Haematopota validicornis, Ricardo, Rec. Ind. Mus., iv, p. 333, pl xvii, fig. 23
(1911).

Although this species is not represented in Dr. Barnes’s collection, the British
Museum (Natural History) possesses a few examples of it from Siam, viz.: 2 99,
precise locality unknown, vi.1906 (W. Palmer); 1 9 (type of H. validicornis, Ric.),
Biserat, 20.x.1901, “‘in spider’s web in jungle’’ (Robinson and Annandale) ; 1 9,
Phrapatoon,* 1907 (Dr. P. G. Woolley). In addition to the foregoing, the National
Collection contains a 9 of H. pachycera from the Federated Malay States (Dr. A. T.
Stanton, presented by the Imperial Bureau of Entomology).

According to the describer of the species, the type of H. pachycera was obtained
in the Laos Pr otectorate, French Indo-China ; Miss Ricardo, however, who examined
the specimen in the Paris Museum, states (Ann. Mag. Nat. Hist. (8) 1, p. 59 (1908) }
that it is from Cambodia.

Haematopota cilipes, Big.
Haematopota cilipes, Bigot, Nouv. Archiv. Mus. Hist. Nat., Paris (3) ii, p. 205

(1893).
According to Bigot (loc. cit., p. 206), the type of this species was collected in
“Laos.” The specimen itself (which the writer has been enabled to examine

through the kindness and courtesy of Baron J. M. R. Surcouf), however, bears labels
stating that it was obtained by M. Pavie in 1886, between Chantabun and
Battambang, both of which localities are in Southern Siam. 4H. cilipes, which is not
included in Dr. Barnes’s collection, is represented in the British Museum (Natural
History) by a solitary 2 from Cambodia, 1909 (John Surcouf, presented by Baron
J. M. R. Surcouf).

* This place-name is printed here and elsewhere as given on labels attached to the specimens
concerned ; extensive search in maps and gazetteers, however, has so far failed to identify this
Siamese locality.

SOME SIAMESE TABANIDAE. 433

Genus Tabanus, Linn.

Hitherto, so far as it has been possible to discover, only three species of
Tabanus—T. rubidus, Wied., T. brunnipennis, Ric., and T. siamensis, Ric.—have
been recorded or described as occurring in Siam. In the following pages the number
of recognised Siamese species of Tabanus is raised to fourteen, namely :—

1. Tabanus barnesi, sp. n. 8. Tabanus striatus, Fabr.

2. ,, migrotectus, Big. ee » virgulatus, sp. n.
os 4 insidiator, sp. Nn. 10. * vubidus, Wied.

4, », praematurus, sp. n. i; » pugnax, sp.n.

Sy » finalis, Walk. 12. » pugrunculus, sp. n.
6. - rubtcundulus, sp. n. 13; i agnosc ibilis, sp. n.
7: - brunnipennis, Ric. 14. 7 stamensis, Ric.

It might be supposed, especially in view of what has been stated above with regard
to the two species of Haematopota mentioned, that one or more of the four species of
Tabanus—T. leucosparsus, T. nigrotectus (Bellardia nigrotecta), T. (Atvlotus)
melanognathus and T. (Atylotus) laotianus—described by Bigot (Nouv. Archiv. Mus.
Hist. Nat., Paris (3) ii, pp. 203-205 (1890) ) from material stated to have been
collected by M. Pavie in the Laos Protectorate,* French Indo-China, would be likely
to occur in Siam. As will be seen below, in the case at least of T°. nigrotectus this
surmise is certainly correct.

Of the previously described species recorded in the following pages, two at
least—T. striatus, Fabr., and 7. rubidus, Wied.—are widely distributed in the
Oriental Region. On the other hand, certain species (7. barnest, sp. n., T.1msidiator,
sp. n., I. praematurus, sp. n.) described below appear to be closely allied to, if not
the representatives of, other forms which occur in the Naga and Lushai Hills, Assam,
so that in their cases the mountain ranges of Upper Burma would seem to form
an effective barrier.

Although the subjoined Key has been tested and found sufficient for the deter-
mination of the extremely limited amount of material at present available to its
author for comparison, infallibility under all circumstances is far from being claimed
for it. Nevertheless, it is hoped that it may prove better than nothing, and may
also serve to stimulate potential collectors of Tabanids in Siam.

Even assuming that no change in the nature of the characters employed in the
Key will be necessitated by the acquisition of further material, it should be noted
that, in the case of a given species shown in the table, these characters are only
sufficient to distinguish it from the other species included in the present synopsis ;
they are not necessarily distinctive as regards additional species of Tabanus, which
may ultimately prove to form part of the Siamese fauna.

It has been necessary to confine the Key to the female sex, since, in the case
of the majority of the species tabulated therein, the male is at present unknown.
Even so, the characters given for Tabanus pugiunculus, sp. n., are perhaps incorrect,
since, as will be seen below (p. 453), it is not absolutely certain that the solitary
female provisionally assigned to that species really belongs to it.

The number in square brackets [ ] after the name of a species indicates the serial
position of the species in the ensuing pages.
Key to the Fourteen Species of Tabanus recorded below (Females only).
1 (2). Wings with first posterior cell closed and petiolate.

(a) Scutellum smoke-grey pollinose and clothed with silvery-white hair,
forming a sharp colour-contrast with remainder of body ; smaller
species, about 15 mm. in length .. sie es barnest, sp n. [1]

*In Kertész’s Cat. Dipterorum, Vol. III, somewhat quaintly styled the Laos Islands
( Ins. Laos”’),

434

MAJOR E. E. AUSTEN.

(b) Scutellum not smoke-grey pollinose nor clothed with silvery-white

13 (18).

14 (15).
15 (14).
16 (17).

17 (16).

18 (13).

hair, entire dorsum of thorax dark mouse-grey, clothed with
blackish hair (abdomen black) ; much larger species, about 20 mm.
inlength .. Br * Sg ss 7 nigrotectus, Big. [2]

. Wings with first posterior cell open.
. Colour of scutellum (and hind margin of scutum) sharply contrasting

with that of dorsum of thorax between bases of wings ; principal
pale markings on dorsum of abdomen in shape of two transverse
bands; small species .. a insidiator, sp. n. [3]

. Colour of scutellum not sharply contrasting with that of thorax

between bases of wings, but either identical or at most somewhat
paler.

. Dorsum of abdomen beyond the base with paler markings confined

to the extreme hind eae: of the second and two following
segments .. a ake .. praematurus, sp. n. [4]

Ni Dorsum of abdomen not so mar ked.
. Dorsum of abdomen beyond the base with paler markings including

or consisting of a series of median triangles or more or less triangular
spots, either separate or in part confluent.

. Last two or last three segments of abdomen presenting a sharp

colour-contrast with remainder ; ae median triangles, large and
broad* ae , stamensis, Ric. [14]

. No colour -contrast “between last two. or last three segments of

abdomen and remainder; ground-colour of dorsum of abdomen
uniform.

1). Median triangles or spots on third and two following abdominal

tergites more or less confluent .. : finalis, Walk. [5]

. Median triangles on second and _ three following abdominal tergites

widely separate .. .. rvubscundulus, sp.n. [6]

. Dorsum of abdomen beyond the base with paler markings including

or consisting of a longitudinal median stripe—not triangles or spots,
although exceptionally the longitudinal median stripe may take the
form of a series of confluent truncate triangles.
Paler markings on dorsum of abdomen consisting solely of a more
or less distinct longitudinal median stripe (no admedian pale
markings—not even a sharply defined, pale spot on each side of
second tergite).
Wings distinctly infuscated (strongly tinged with sepia)
pugnax, sp. 0 site
Wings hyaline.
Wing-stigma conspicuous ; front relatively rather broad (4 to 4}
times as long as its breadth between the lower inner angles of the ey es)
pugiunculus, sp.n. [12]
Wing-stigma inconspicuous ; front narrow (about seven times as long
as its breadth between the lower inner angles of the eyes) .
agnoscibilis, sp. n. _ [13]
Paler markings on dorsum of abdomen, in addition to the longi-
tudinal median stripe, consisting of an admedian stripe or an
admedian longitudinal series of spots on each side, or at least of
a sharply defined pale spot on each side of second segment.

* Owing to the completely denuded condition of the type and paratype of T. siamensis, Ric.
(see below, p. 455), the only specimens of this species at present available, the absolute accuracy
of this statement as to the triangles cannot be guaranteed, though the ground-colour presents
indications which seem to warrant it.

SOME SIAMESE TABANIDAE. 435

19 (20). Admedian paler markings on dorsum of abdomen practically confined
to second segment, which bears a pair of admedian spots (similar
spots, if present on oe of following tergites, usually much less
cistinct)*, 4). .. brunnipennis} Ric. [7]
20 (19). Admedian paler markings on dorsum of abdomen not confined to a
pair of spots on second segment, but much more extensive (see 18
above).
21 (22). Median longitudinal stripe on second abdominal tergite more or
less obsolete, at least much less distinct and well developed than
on following tergite.. ‘e striatus, Fabr. [8]
22 (21). Median longitudinal stripe on "second abdominal tergite not obsolete,
in any case just as distinct and well developed as that on following

tergite.
23 (24). Frontal callus blackish-brown ; expanded portion of third segment
of antenna cinnamon-rufous.. .. virgulatus, sp.n. [9]

24 (23). Frontal callus chestnut-brown or russet- coloured ; expanded portion
of third segment of antenna mainly or entirely blackish or blackish-
brown ae sis i is a . rubtdus, Wied. [10]

1. Tabanus barnesi, sp. n. (fig. 1).

2. Length (one specimen), 15 mm. ; width of head, just under 5 mm. ; width of
front at vertex, 0-6 mm. ; between lower inner angles of eyes, 0-4 mm. ; length of
wing, 15 mm.

Fig. 1. Head of Tabanus barnesi, Austen, sp. n., ©) ; a, front view; 8, profile.

Dorsum of thorax, except scutellum, olivaceous-black,* scutellum, except extreme
base, smoke-grey pollinose, clothed above with silvery white hair ; abdomen black, first
five (visible) tergites each with a transversely elongate patch of short, whitish, appressed
hairs in each posterior angle; wings with costal cells light cinnamon-brown, and a
mummy-brown transverse band before distal extremity ; first posterior cell closed about
0:5 mm. from wing margin.

+ Or names and illustrations of colours used for descriptive purposes in the present paper
see Ridgway, ‘‘ Color Standards and Color Nomenclature ’’ (Washington, D.C. Published by
the Author, 1912).

436 MAJOR E. E. AUSTEN.

Head: Subcallus drab, front, face and jowls greyish-olive pollinose, face and
jowls clothed with fine dark-brown or blackish-brown hair, occiput pale smoke-grey
pollinose, clothed with fine hair of similar colour, occipital margins fringed posteriorly
with short, glistening yellowish hair, except behind vertex, where the somewhat
longer hairs are blackish ; front in 2 narrow, somewhat wider above than below,
clothed with minute, appressed, glistening yellowish hairs, vertex also bearing short
blackish hairs, in typical specimen with a median, dark, somewhat shining, elongate
mark, but showing no trace of an ocellar tubercle; frontal callus, blackish-brown,
longitudinally elliptical in outline, and continuous above with a median rib-like
extension of same colour, which, in case of typical specimen, reaches a point well
above midway between lower end of fvont and hind margin of vertex; eyes bare,
without recognisable bands in dried condition ; palpi, drab, proximal segment clothed
with fine dark brown or blackish-brown hair, distal segment in 9 elongate, bluntly
pointed at distal extremity, and clothed on outer surface with minute, appressed,
black hairs, among which, especially at base above, a number of minute, glistening
yellowish hairs is visible; first and second segments of amtennae cinnamon-buff
(pinkish-buff when viewed from above), clothed with minute black hairs, first segment
partly embracing second, its upper distal angle being prominent, upper distal angle
of second segment considerably produced, third segment cinnamon (infuscated “at
extreme tip in case of type), fairly deep at base, with a prominent though not elongate
angle on upper border, after which expanded portion is somewhat elongate, the part
of expanded portion beyond the angle being about twice the length of the part from
base to angle inclusive, total length of annulate portion of third segment about half
that of expanded portion. Thorax: Dorsum (except anterior border, humeral calli,
and scutellum, with exception of its black base) clothed with black hair, which is
thus present on base of scutellum also; humeral calli clothed with greyish-white
hair, upper portion of anterior border of dorsum sparsely clothed with minute
appressed, glistening yellowish hairs, a few similar hairs also present among the
adjacent black hair; pleurae clothed partly with black, partly with brownish hair ;
anterior surface of thorax, next to occiput, olive-grey pollinose ; pleurae and pectus
dark brownish olive-grey pollinose, a thin coating of similar character and colour
also present on dorsum of scutum, at least anteriorly, although, at any rate in a
partially rubbed specimen, dorsum appears somewhat shining. Abdomen: Dorsum
somewhat shining ; whitish hairs forming lateral patches on dorsum fine and some-
what thinly set, lateral patches on first (visible) segment confined to the actual
posterior angles, on following segments corresponding patches are much larger, each
patch occupying on lateral margin at least posterior half of the segment, diminishing
in depth towards middle line while remaining in contact with hind margin of tergite,
and extending over about one-fourth (on fifth tergite about one-third) of the width
of the segment; sixth tergite with a few scattered whitish hairs near each lateral
extremity ; dorsum, except as stated, clothed with short, appressed black hair ;
venter brownish-black, moderately shining, clothed with short, appressed black hair,
among which on hind borders of second and two following sternites, and also scat-
tered over surface of fourth sternite, are short or minute, glistening whitish hairs.
Wings: Veins mainly dark brown, second longitudinal vein partly, third longitudinal
at base, and anterior transverse vein tawny-olive ; anterior branch of third vein
(at least in type) with a small appendix (in case of type, more noticeable in right wing
than in left) ; sé¢gma, tawny-olive, elongate and inconspicuous ; first basal cell faintly
tinged with sepia ; mummy-brown transverse band wider on costa than towards
hind margin, and dying away before reaching latter, but extending into fourth
posterior cell; on costa, band extends from a point about half-way along lower
margin of stigma to end of first submarginal cell; thence, distal margin of band
runs obliquely backwards, showing a more or less pronounced indentation in second
submarginal cell ; proximal margin of band straighter, running across wing practi-
cally at right angles to longitudinal axis of latter, and including distal extremity

SOME SIAMESE TABANIDAE. 437

of discal cell (in the case of the type, in practically every cell crossed by the band
the latter is interrupted by a large hyaline streak, so that the band itself is composed
of mummy-brown borders to the veins, but this condition is doubtless due to individual
aberration). Squamae, deep mouse-grey, scantily fringed with fine yellowish hair ;
borders of squamae mummy-brown, outer edges ivory-yellow. Halteres, cream-buff,
stalks and under surface of knobs more orlessmummy-brown. Legs: entirely black
and clothed exclusively with black hair; front tibiae not thickened ; front tarsi
in © not conspicuously expanded, but with distal angles of penultimate segment
considerably produced.

Chiengmai, 10.v.1921. With reference to the holotype of this species, Dr. M. E.
Barnes, in whose honour the author has much pleasure in naming it, writes as

follows :—‘‘ Caught at about 5.0 o’clock p.m., attacking me while I was seated on
the verandah of my house ; this is the only specimen that I have seen.”’

In addition to the foregoing specimen, the British Museum (Natural History)
also possesses an old, considerably damaged, and much faded @ of this species, taken
upwards of fifty years ago at Chantabun, S. Siam (— Mouwhot), and formerly in
the collection of the late W. W. Saunders. While certain differences from the specimen
selected as the type are noticeable, none of the points as to which divergence appears
can be regarded as more than varietal, and in some respects, as in the condition of the
transverse band on the wing, the example from Chantabun is probably the more
truly typical of the two. In the Chantabun Q the rib-like upward extension of the
frontal callus is concealed by the pollinose covering of the front, from a point a little
below the middle of the latter; so far as can be seen, there is no trace of lateral
patches of whitish hair on the dorsum of the abdomen; the anterior branch of
the third vein is without even a vestige of an appendix; and the mummy-brown
transverse band on the wing is not interrupted by hyaline streaks.

Tabanus barnesi resembles T. (Atylotus) nephodes, Bigot, of which the type, from
the Naga Hills, Assam (Captain Butler), and a second 9, from Sibsagar, Assam,
are now in the National Collection. While agreeing with the species in question
in its wing-marking, and in the first posterior cell being closed at some distance from
the wing margin, 7. barnesi is, however, distinguishable (in the 9 sex) inter alia by its
smaller size; by the frontal callus being, if anything, somewhat larger and more
clearly differentiated from its rib-like upward extension ; by the jowls, pleurae and
lower part of the face being clothed with dark brown or black, instead of with whitish
hair; by the dorsal surface of the scutellum, except at the base, being smoke-grey
pollinose, and clothed with glistening silvery-white hair; by the total absence of
median, white-haired triangles on the dorsal surface of the abdomen ; and by the
appendix to the anterior branch of the third vein, which is long in T. nephodes, being
vestigial or absent.

2. Tabanus nigrotectus, Big.
Bellardia nigrotecta, Bigot, Nouv. Archiv. Mus. Hist. Nat., Paris (3). i,
p. 204 (1890).

Though stated by Bigot to be from “ Laos,” the type of this species, which
Baron Surcouf has most courteously sent to the writer for examination, bears labels
identical with those attached to that of Haematopota cilipes, Bigot (see above, p. 432).
T. nigrotectus therefore occurs in Southern Siam. The species was not among those
of which examples were forwarded by Dr. Barnes, but is represented in the National
Collection by a Q from Cambodia, 1909 (John Surcouf, presented by Baron J. M. R.
Surcouf).

3. Tabanus insidiator, sp. n. (fig. 2).

©.—Length (five specimens), 10 to 10:6 mm.; width of head just under
4to4-5mm.; width of front at vertex, 0-6 mm. ; length of wing, 9:4 to 10 mm.

438 MAJOR E. E. AUSTEN.

Pretty little species, with, in 9, two frontal calli, bare eyes having apparently a single
transverse purple band on level of lower callus, brightly marked body, and parti-coloured
legs. —Pleurae light greyish-olive pollinose, scutellum pale olive-buff pollinose, both
clothed conspicuously with pale yellowish hair; dorsum of abdomen mainly chestnut
brown at base, then black or blackish-brown, third and fourth (visible) tergites each with
a very conspicuous, pale (olive-buff pollinose) hind border, clothed with appressed
glistening yellowish hair, and in each case expanded in middle into a wide, low triangle.

Head: Subcallus (in all five specimens available for comparison) shining russet-
brown or mummy-brown, sometimes with a small, blackish-brown, median spot above
bases of antennae ; front greyish-olive pollinose, darker above, face and jowls pallid
neutral grey pollinose, clothed with yellowish hair, upper extremities of sides of face
immediately below subcallus usually mummy-brown pollinose, occiput smoke-grey
pollinose, thinly clothed with pale yellowish hair, posterior orbits on each side of
vertex fringed behind with minute black hairs mixed with minute, glistening yellowish
hairs, posterior orbits elsewhere fringed with minute, glistening yellowish hairs ;
front, in 2 of moderate width, somewhat narrower below than above, and in length
equal to from five to six times its breadth between lower inner angles of eyes, clothed

Vig. 2. Head of Tabanus insidiator, Austen, sp.n., S) ; a, front view; 0, profile.

above with minute black hairs which are longer on vertex, where there is an ill-
defined dusky patch but no trace of an ocellar tubercle ; frontal calli (see fig. 2a),
blackish-brown (lower callus sometimes reddish-brown), lower callus shield-shaped
or quadrate, its lower margin (except point if shield-shaped) a little above level of
lower inner angles of eyes, upper callus elongate quadrate or roughly elliptical,
sometimes divided in middle line above, and in some specimens drawn out into a
point below, or with a lower median extension which may or may not reach lower
callus ; palpi, neutral grey or deep neutral grey on outer, pallid neutral grey on inner
surface, distal segment bluntly acuminate, moderately swollen at base, proximal seg-
ment, base and proximal two-thirds of lower margin of distal segment clothed with
yellowish hair, outer surface of distal segment clothed elsewhere with minute, ap-
pressed black hairs ; first two segments of antennae pale ochraceous-tawny, expanded
portion of third segment tawny or russet, annulate portion cinnamon-brown, second
segment and upper portion of first segment clothed with minute black hairs, first
segment clothed below with yellowish hair and considerably swollen distally, having
its upper distal angle moderately produced and partly embracing second segment,
upper distal angle of latter small and not conspicuously elongate, expanded. portion of
third segment of moderate depth, varying from one-fourth as long again to nearly
twice as long as annulate portion, angle on upper margin near base conspicuous

SOME SIAMESE TABANIDAE. 439

but not produced. Thorax: Dorsum—except fore border, scutellum, humeral and
post-alar calli, swelling occupying depression at each end of transverse suture, a
small area in front of suture immediately above each of these swellings, and extreme
hind margin of scutum—blackish-brown, and clothed with similarly coloured hair ;
anterior surface and fore border of dorsum smoke-grey or light greyish-olive pollinose,
the pollinose area extending on each side to transverse suture, and clothed with
appressed, glistening yellowish hair; swelling in depression at each end of transverse
suture, and area immediately above it in front of suture, pinkish-buff or cinnamon-
buff, covered with olive-buff pollen and clothed with longer bright yellowish or ochreous
hair, swellings at ends of suture also with a certain number of black or blackish hairs
below ; humeral and post-alar calli, and extreme hind margin of scutum pale smoke-
grey pollinose, and clothed with yellowish hair; pectus agreeing with pleurae in
coloration and hairy covering. Abdomen: First and second (visible) tergites mainly
chestnut-brown, remainder of dorsum, except as already or subsequently described,
black or blackish-brown ; first (visible) tergite blackish-brown in centre; second
tergite with a, not sharply defined, median triangular patch and a blotch on each side
(not extending into posterior angle) of same colour ; lateral extremities of first (visible)
tergite, and posterior angles of second tergite, olive-buff pollinose and clothed with
yellowish hair ; hind margins of fifth and sixth tergites (sometimes that of seventh
tergite also) narrowly olive-buff pollinose, and clothed, at least in part, with hair of
same kind as that on hind borders of third and fourth tergites ; dorsum, except as
stated, clothed with short appressed black hair; venter at base, as far as hind margin
of second segment, pale cinnamon or light pinkish-cinnamon pollinose, clothed with
yellowish hair ; remainder of venter agreeing with dorsum in coloration, markings and
hairy covering, except that pale hind borders on third and fourth segments, though of
same depth as above, do not exhibit median, triangular expansions. Wings strongly
tinged with mouse-grey ; veins mummy-brown or cinnamon-brown, anterior branch
of third longitudinal vein almost rectangular at base, and with an appendix which
varies in size in different individuals, and occasionally is entirely wanting ; stigma
tawny-olive, usually fairly well defined and conspicuous. Sguamae dusky, borders
dark brown, fringed with fine, short pale hair. Halteres dark brown or blackish-brown.
Legs: Coxae and trochanters light greyish-olive pollinose, clothed with yellowish hair,
hind coxae, at least in front, usually clothed mainly with black hair ; femora black,
usually more or less greyish pollinose, at least beneath, front femora clothed with black
hair, those of middle and hind legs clothed partly with black, partly with yellowish
hair; tibiae cream-buff, approximately distal third of front tibiae, and tips of middle
and hind tibiae black or blackish-brown and clothed with black hair, cream-buff area
of tibiae clothed with fine, glistening silvery hair, with which black hairs (sometimes
especially numerous on extensor surface of hind tibiae) may be intermixed ; tarsi
black, clothed with black hair, third and fourth segments of front tarsi in 9 only slightly
expanded.

Holotype and four paratypes caught on Doi Chom Chang, near Chiengmai,
N. Siam, alt. 5,000 ft., 16.iv.1921. With reference to these specimens, Dr. Barnes
writes : ‘“‘ Taken while attacking me as I was walking through some jungle. This
fly, the bite of which is quite painful, is very common in the jungle on the mountain.”

The species just described, although strongly resembling and closely allied to
Tabanus (Atylotus) leucocnematus, Big., is, in the 2 sex, distinguishable therefrom
inter alia by the frontal calli being wider apart; by the expanded portion of the
third segment of the antenna being shorter and also deeper at the base ; by the short
appressed hair clothing the scutellum, post-alar calli, hind margin of the scutum,
and hind borders of the third and fourth abdominal tergites being much paler; by
the median triangular expansions of the abdominal bands in question being smaller ;
by the black or blackish-brown tips to the front tibiae being at least twice as deep ;
and by the wings being somewhat paler, more elongate, more acuminate, and having
a more clearly marked stigma.

440 MAJOR E. E. AUSTEN.

It may be added that, although the precise origin of the type of T. leucocnematus,
Big. (now in the National Collection), is unknown, since Bigot (Mém. Soc. Zool.,
France, v., p. 657 (1892)) records it merely as “ Indes,” the specimen may well have
been taken in Assam. At any rate, the British Museum (Natural History) possesses
a second ° of T. leucocnematus, Big., which was captured at Kolasil, Lushai Hills,
Assam, 30.v1i.1904, by Captain E. C. Macleod.

4. Tabanus praematurus, sp. n. (fig. 3).

°.—Length (onespecimen) 16-2 mm. ; width of head, 6 mm. ; width of front at
vertex, Just under 1 mm., between lower inner angles of eyes, 0-4 mm.; length
of wing, 15 mm.

Eves bare, apparently with two (three?) purple bands ; a single narrow, elongate,
frontal callus in 2°; dorsum of thorax (scutum) tawny-olive pollinose, front and hind
borders (including humeral and post-alar call1) and scutellum paler ; abdomen, except
first (visible) segment, warm blackish-brown (last three segments darker), hind margins
of second to fourth tergites inclusive, and of fifth sternite narrowly whitish, and clothed

Fig. 3. Head of Tabanus praematurus, Austen, sp. n., ) ; a, front view; 8, profile.

(except middle third in case of fifth sternite) with appressed yellowish or silvery-white
hair, second to fourth sternites inclusive with deeper whitish-grey pollinose hind borders,
especially towards the sides, clothed with appressed, glistening, silvery-white haw ;
wings with base, costal and basal cells tinged with tawny-olive, elsewhere, with exception
of a larger or smaller hyaline fleck in almost every cell, suffused with sepia ; legs, except
coxae, trochanters, extreme lips of femora and bases of middle femora, uniformly blackish-
brown or black.

Head: Subcallus, upper part of sides of face and lower half of front cinnamon-
brown pollinose, upper half of front darker (mummy-brown), jowls and lower part
of face buff-yellow pollinose, occiput ochraceous-buff pollinose, posterior orbits
smoke-grey ; upper part of sides of face clothed with dark brown hair, jowls and
occiput clothed with buff-yellow hair, hind margin of occiput sparsely fringed above
on each side of front with glistening hair of somewhat deeper tint ; from in 9 narrow

SOME SIAMESE TABANIDAE. 44]

at its lower extremity, but from level of upper end of main portion of callus to vertex
increasing considerably in width (see fig. 3a), sparsely clothed below and on lateral
margins with appressed, glistening ochreous hairs, and above with short, erect
blackish hair, vertex with a depressed, shining black median area shaped something
like a truncate triangle, but with no trace of an ocellar tubercle ; frontal callus blackish-
brown, narrow and elongate, its upper extremity produced into a stout, linear ex-
tension, slightly longer than main portion of callus, and reaching to a point a little
above midway between level of lower inner angles of eyes and hind margin of vertex ;
proximal segment of palpi clothed below with buff-yellow hair and on upper border
of outer side with black hair, distal segment in 2 acuminate and rather narrow,
mouse-grey on outer side except at extreme base, inner side and extreme base of
outer cinnamon-drab, outer surface clothed with minute, appressed black hairs,
a few glistening buff-yellow hairs at base below ; first and second segments of antennae
light fuscous or olive-brown, clothed with black hair, first segment broadening from
base to tip, with its upper distal extremity partially enveloping second segment,
upper distal angle of latter strongly produced, third segment russet, expanded portion
slightly longer than annulate portion, fairly deep at base, with angle on upper border
strongly produced upwards and forwards (see fig. 3). Thorax: Pleurae dark olive-
buff pollinose, scutellum somewhat paler pollinose, dorsum of scutum (in partially
rubbed specimen) with more or less distinct traces of four ill-defined, dark brown,
longitudinal stripes ; dorsum of scutum clothed with fine, erect, glistening honey-
yellow hair, pleurae and dorsal surface of scutellum clothed with similar hair of a
paler (yellowish) tint. Abdomen: Tergite of first (visible) segment isabella-coloured,
clothed with fine, short, appressed, glistening honey-yellow hair ; remaining tergites
clothed for most part with short, appressed, black hair, with which on central portion
of second tergite honey-yellow hair like that on first tergite is freely mingled ; hair
clothing whitish pollinose hind margins of second to fourth tergites mainly silvery-
white, but some glistening honey-yellow or ochreous hairs also present on or just
in front of hind margins of second and third tergites, except at sides; venter at
extreme base greyish-olive or neutral grey pollinose ; sternite of (nominal) second
segment, on each side in front of hind border, clothed for most part with minute,
appressed, glistening yellowish hairs; seventh segment clothed above and below
with longer black hair ; sternites of (nominal) second and following segments, except
as already stated, clothed with minute, appressed black hair. Wings: Stigma
ochraceous-tawny, elongate and tapering ; costa, and main stem of third longitudinal
vein except at base, mummy-brown, veins otherwise for most part cinnamon-brown
or amber-brown ; in addition to usual hyaline streak in base of marginal cell, before
stigma, and usual hyaline spot on fourth longitudinal vein, immediately before origin
of anterior basal transverse vein, all cells, except costal, basal and anal cells, show—
at least in typical specimen—a more or less distinct hvaline mark, varying in size
and shape in the different cells. Squwamae pale isabella-coloured (margins lighter or
darker according to incidence of light), fringed with pale yellowish hair. Halteres
cream-buff, base of knobs (in dried condition) cinnamon-brown. Legs: Coxae dark
olive-buff pollinose, clothed with yellowish hair like that on pleurae ; front and hind
trochanters mouse-grey, hind pair clothed partly with yellowish, partly with dark
brown hair, trochanters of middle legs cinnamon, with darker markings ; middle
femora cinnamon at base, extreme tips of all femora cinnamon-buff ; tibiae, tarsi,
and femora for most part clothed with black hair, posterior surface of middle femora,
except at tips, clothed with long and fine yellowish hair, glistening ochreous hairs
also present at base of middle femora in front, and at base of hind femora above and
below, front femora fringed posteriorly with fairly long black hair ; front tibiae not
thickened, second and following segments of front tarsi, especially fourth segment,
considerably expanded in 2; claws long, entirely black ; pulvilli ochraceous-buft.

Doi Chom Chang, near Chiengmai, alt. 5,500 ft., 15.1v.1921. Writing of the holo-
type of this species, Dr. Barnes says: ‘‘ This specimen was caught while attacking

442 MAJOR E. E. AUSTEN,

me at dawn in my cottage. On three mornings I was awakened about day-break
by attacks by one or more of these flies, which I did not see on any other occasion.”’

The species described above is allied to Tabanus manipurensis, Ric., represented
in the British Museum (Natural History), by the holotype 9 from Ukhrul, Manipur, alt.
6,400 ft. (Rev. W. Pettigrew). Although not unlike 7. manipurensis in general
appearance, 7. praematurus is distinguishable, inter alia, in the 2 sex by the front
being narrower below ; by the expanded portion of the third segment of the antenna
being much shorter, and having the angle on its upper border much more strongly
developed ; by the tibiae being uniformly blackish-brown or black instead of con-

spicuously creamy-white or cream-buff except at the distal extremity and extreme ©

base ; and by the greater breadth of the last three joints of the front tarsi.

5. Tabanus finalis, Walk.
Tabanus apicalis, Walk., List. Dipt. Ins. in coll. Brit. Mus., i, p. 176 (1848).—
Nomen bis lectum.

Tabanus finalis, Walk., op. cit., v. Suppl. 1, p. 258 (1854).

Although this species is not included in Dr. Barnes’s collection, the British Museum
(Natural History) possesses a single 9 of it taken in Siam (precise locality uncertain),
3.11.1914, “ at light ’”’ (K. G. Gairdner). The provenance of the type (a 3) is unknown.

Tabanus finalis is a fairly large (about 18 mm. long), blackish-brown insect of
striking appearance, recognisable by the presence of a large, cream-buff median
spot on each abdominal tergite from the third to the fifth, inclusive ; on the second
(visible) abdominal tergite there are three small pale spots in a transverse row, of
which the middle one is drab-coloured, while the two lateral ones are smoke-grey ;

the wings are infuscated, and the veins in the central area to a greater or less extent |

have brownish borders.

6. Tabanus rubicunduius, sp. n. (fig. 4).

2.—Length (two specimens) 18 to 19 mm.; width of head, 5-6 to6 mm. ; width |
of front at vertex 0-75 mm., between lower inner angles of eyes 0-4 mm., length of |

wing 16-25 to 17-2 mm.

Fig. 4. Head of Tabanus rubicundulus, Austen, sp. n., 2 ; a, front view ; 0, profile.

Eyes bare; a single elongate frontal callus in 2; ground-colour of dorsum of body |
chestnut-brown ; dorsum of abdomen with a median series of conspicuous, light buff |

SOME SIAMESE TABANIDAE. 443

triangles ; anterior branch of third longitudinal vein in wing with a well marked
recurrent appendix.

Head: Front and subcallus olive-buff pollinose, upper half of front appearing darker
when seen from certain angles (looking almost mummy-brown when viewed at a
low angle in an antero-posterior direction) ; face, jowls and occiput pale smoke-grey
pollinose, clothed with whitish hair, upper extremities of sides of face faintly
suffused with sepia and clothed with dusky or blackish hair, hind margin of upper
part of posterior orbits with an inconspicuous fringe of very short black hair, front
in 2 clothed with minute dusky hairs; front in 2 narrow, diminishing in width
from above downwards, and about eight or between eight and nine times as long as its
breadth between lower inner angles of eyes, ocellar tubercle wanting ; frontal callus
(see fig. 4a), dark chestnut-brown, vertically elongate (roughly elliptical—its base
just above lower inner angles of eyes), exhibiting a median impressed line, and
continued above into a linear prolongation; palpi in 2 drab-coloured, proximal
segment clothed with whitish hair, distal segment narrow and elongate, blunt at
tip and but little thicker at base, clothed on outer surface with minute, appressed
black hairs ; antennae, russet, first and second segments clothed with minute black
hairs (with yellowish hairs on lower margin of their distal extremities), first segment
somewhat greyish pollinose above, strongly swollen distally, its upper distal angle
produced and partly embracing second segment, upper distal angle of latter con-
siderably elongate, expanded portion of third segment in @ of considerable depth
at base, not elongate and having a prominent, blunt angle half-way along its upper
margin, annulate portion of third segment (dark chestnut-brown in case of type)
about one-fourth shorter than expanded portion. Thorax: Dorsum, including
scutellum, thinly, pleurae and pectus densely smoke-grey pollinose; dorsum,
including scutellum, clothed with semi-erect black hair, mixed with fine, appressed
or recumbent, ochreous or yellowish hair, a tuft of whitish hair also present above
base of each wing and on post-alar calli ; swelling occupying depression at each end
of transverse suture, as well as post-alar, calli in part and a small area on each side
behind and above tuft of whitish hair above base of wing clothed with black hair ;
pleurae and pectus clothed with whitish hair. Abdomen: tergites of second to fifth
segments inclusive each with a light buff, median triangle resting on its hind margin,
the triangle in each case being formed by a patch of appressed, glistening straw-
yellow hairs clothing a light buff pollinose triangular area; in case of holotype and
paratype, triangle on second segment is isosceles, and its apex extends somewhat
beyond middle of length of segment, while remaining triangles are equilateral and,
except in case of triangle on fifth tergite, do not extend as far as middle of their
respective segments ; lateral extremities of first (visible) tergite pallid neutral grey
pollinose, ground colour of lateral extremities, or at least posterior angles of following
five tergites ochraceous-tawny ; lateral extremities of first (visible) and three following
tergites clothed with whitish or silvery-white hair ; a small, triangular patch of ap-
pressed, glistening straw-yellow hairs on hind margin of first (visible) tergite in
middle line, sixth tergite with a more or less distinct, median triangular patch of
similar hair, interspersed with minute black hairs, the patch resting on hind margin
and showing individual variation in size, being sometimes very small, in other cases
considerably larger, so that its apex almost reaches hind margin of preceding
segment ; owing to absence of a light buff, pollinose ground beneath the median
patches of appressed straw-yellow hairs on the first and sixth tergites, these patches
are somewhat inconspicuous and do not appear as clearly defined triangles, con-
trasting sharply with surrounding area, as in the case of the median markings on the
four intervening segments; fourth and fifth tergites with a few glistening straw-
yellow hairs on their posterior angles; lateral extremities of last three, and hind
margins of last two tergites clothed with black hair of moderate length ; dorsum,
except as stated, clothed with minute, appressed black hairs; venter russet, hind
margins (or at least their lateral extremities) of second to fifth sternites, inclusive,

(5296) 2K

444 MAJOR E. E. AUSTEN.

narrowly light buff or cream-buff, entire ventral surface with a thin, pale drab-
grey, pollinose covering, which, however, when venter is viewed from behind at a
low angle, allows a large, quadrate, not sharply defined, dark median blotch to be
seen on each segment from second to sixth inclusive ; venter clothed mainly with
fine, minute black or blackish hair, seventh sternite clothed as usual with coarse,
erect black hairs, lateral thirds of second sternite, lateral fourths of two following
sternites except anteriorly, and posterior angles of fifth sternite clothed with
glistening, appressed silvery white or yellowish hair, which is longer at lateral
extremities of hind margins than elsewhere ; entire hind margins of second to fourth
(or perhaps, second to fifth) sternites inclusive, sometimes clothed with similar
hair. Wings sepia-coloured (in life probably considerably darker), an ill-defined,
slightly paler area in second submarginal cell; veins lighter or darker mummy-
brown, anterior branch of third longitudinal vein forming a right angle with main
stem, then bent at an obtuse angle, appendix (at least in case of type and paratype)
between 0-4 and 0-5 mm. in length; sé#igma sepia-coloured or dark tawny-olive,
narrow, elongate and tapering. Sguamae sepia-coloured, borders mummy-brown,
fringed with short pale hair; antisquamae fringed with longer whitish hair. Halteres
cinnamon-brown, stalks, except distal extremities, and tips of knobs paler (light
ochraceous-buff). Legs: coxae drab-grey pollinose, front coxae and outer surfaces
of middle and hind pairs clothed with fine whitish or yellowish-white hair, lower
surfaces of middle and hind coxae, and also part of lower portion of outer surface
of middle coxae, clothed with black or blackish hair ; femora russet-coloured, darker
(more or less blackish-brown) above (at least in case of front legs), upper surfaces of
femora clothed mainly with short, appressed black hair, mixed, at least in case of
hind pair, with glistening ochreous or yellowish hairs, hind femora also with a tuft
of whitish hair at base above ; posterior surfaces of front and middle femora clothed
with longer whitish hair (mixed above with black or blackish hair in case of front
femora), hind femora, with longer whitish or yellowish-white hair below, and with
whitish or ochreous hair on lower portion of anterior surface; front tibiae chocolate-
brown or clove-brown, and clothed with minute, appressed black hairs, base on outer
side paler (indistinctly russet), clothed with minute, glistening, appressed Naples
vellow hairs; middle and hind tibiae russet-brown or russet, paler at base, clothed
mainly with minute, appressed black hairs, proximal portion of inner surface in
case of middle tibiae, and of outer surface in that of hind pair clothed with glistening,
appressed, ochreous or yellowish hair, outer edges of extensor surfaces of hind tibiae
fringed with longer black hair, flexor surfaces of hind tibiae sometimes largely clothed
with appressed, glistening ochreous hairs; front tarsi black, third and following
segments strongly expanded in 2; middle tarsi blackish-brown, likewise with last
three segments somewhat expanded; hind tarsi with first segment and proximal
two-thirds of second segment russet, otherwise dark brown ; all tarsi clothed above
with minute black hairs.

S. Sram: Chantabun (— Mouwhot :—ex coll. the late W. W. Saunders).

In general appearance, as also in the markings on the dorsal surface of the
abdomen, and in the narrowness of the front in the 9, Tabanus rubtcundulus resembles
T. indianus, Ric. (Rec. Ind. Mus., Calcutta, iv., p. 175 (1911) ), which, originally
taken in India (North Kanara, Bombay Presidency), is also found in Hong Kong
and Formosa. The new species described above may, however, at any rate in the
2 sex, be distinguished from the one in question by its differently coloured legs (the
femora being paler, and the proximal halves or three-fifths of the front tibiae not
being cream-coloured), and by the presence of a well-developed appendix to the
anterior branch of the third longitudinal vein.

7. Tabanus brunnipennis, Ric.
Tabanus brunnipennis, Ricardo, Rec. Ind. Mus., Calcutta, iv, p. 160 (1911).

SOME SIAMESE TABANIDAE. 445

Of this species, the type of which is from India (North Kanara), the British
Museum (Natural History) possesses a 9 from Bangkok, collected in 1898 by Major
S. S. Flower, O.B.E. Apart from this specimen and the typical series, the only
examples of the species as yet contained in the National Collection are from South
Malabar (South-west India, a little to the south of the locality where the type and
paratypes were obtained).

8. Tabanus striatus, Fabr.
Tabanus striatus, Fabricius, Entomologia Systematica, iv, p. 371 (1794).
Tabanus partitus, Walk., Journ. Proc. Linn. Soc., i, p. 9 (1857).

One 3, Bangkok, 30.viii.1921, caught in donor’s house at night, attracted by
electric light ; only specimen seen: one 9, Chiengmai, 7.v.1921, on donor’s verandah
at dusk—a solitary specimen, which did not attack. Two 99 of this species from
Bangkok, taken respectively in 1907 (Dr. P. G. Woolley) and November 1919 (F. af
Godfrey), were previously included in the Museum collection.

The distribution of 7. striatus is very wide, its range, as shown by material in the
National Collection, extending right across the Oriental Region, from North-western
and Western India (Kohat and Bombay) to Hong Kong and the Philippine Islands,
and including the Federated Malay States, Singapore, Sumatra and Java.

According to Mitzmain,* who studied the bionomics of this species in the Philip-
pines, and also in the same year (1913) demonstrated experimentally its capacity to
act as a mechanical transmitter of surra,} 7. striatus is ‘‘ the most prevalent horsefly ”
in the Philippine Archipelago, where, although also attacking cattle and horses, it
appears to prey by preference on the carabao (buffalo). As regards human beings,
the author referred to writes: ‘‘ During over two years of personal observation, this
fly has never been known to annoy man in the Philippine Islands.”’

In the dried (pinned) condition, at any rate, apart from their usually darker appear-
ance, specimens of T°. striatus, especially when the median abdominal stripe is not
obliterated on the second tergite, are sometimes liable to be mistaken for examples
of T. tenens, Walk., of which T. hilaris, Walk. (Insecta Saundersiana, i, Diptera. pist
p. 49 (1850)), and 7. megalops, Walk. (List Dipt. Ins. in coll. Brit. Mus., v, Suppl. i,
p. 247 (1854)) aresvnonyms. So far as it is at present possible to judge, the range of
7. tenens is much more restricted than that of T. striatus, though the evidence afforded
by the series of specimens in the British Museum (Natural History) shows that this
species is common in parts of India (e.g. Madras), and also occurs in Ceylon and Java.
In the 3 sex, excluding differences due to colour, T. striatus, Fabr., is distinguishable
from 7. tenens, Walk., by the distal margin of the penultimate segment of the front
tarsi being more deeply notched. Inits typical form, as represented by specimens from
Hong Kong, the 9 of 7. striatus may be distinguished from that of 7. tenens, inter alia,
by the shape of the lower frontal callus, which is less elongate, 7.e., broader in pro-
portion toits length ; by the shape of the third and fourth segments of the front tarsi,
the sides of these segments when the front legs are viewed from above appearing
straighter ; and by the legs in general being darker—the femora infuscated, and the
front tarsi deep black instead of ferruginous or brownish.

Miss G. Ricardo, in her ‘“ Revision of the Species of Tabanus from the Oriental
Region,’’§ involves the identity and synonymy of Tabanus striatus in hopeless
confusion, since 7. tenens, Walk., and T. megalops, Walk., are included among the

* Cf. M. B. Mitzmain, ‘‘ The Biology of Tabanus striatus Fabricus [sic], the Horsefly of the
Philippines ”’ : Philippine Journ. Sci., viii, no. 3, Sec. B, Tropical Medicine, pp.197—221, pls. I-VII
(June 1913).

+ Cf. M. B. Mitzmain, ‘“‘ The Mechanical Transmission of Surra by Tabanus striatus Fabricus ”’
[sic]: ibid., pp. 223-229 (June 1913).

§ Cf. G. Ricardo, “‘ A Revision of the Species of Tabanus from the Oriental Region, including
Notes on Species from Surrounding Countries ’’: Rec. Ind. Mus., iv, pp. 150, 153 (1911).

(5296) DANNS

446 MAJOR E. E. AUSTEN.

synonyms of T. striatus, Fabr., while T. hilaris, Walk., is treated as a valid species.
Curiously enough, in writing of T. hilaris Miss Ricardo says ( loc. cit., p. 153): “A
species distinguished from T. striatus, F., by the short median stripe of abdomen,
which does not begin till the ¢rivd segment and by the shorter lateral stripes which
usually terminate on the third or fourth segment.’’ The author in question thus
appears to regard as distinctive in comparison with T. striatus, Fabr., characters.
which are actually among those given by Fabricius himself as diagnostic of the latter
species.t

It may be added that Tabanus sinicus, Walk. (List. Dipt. Ins. in coll. Brit. Mus...
i, p. 163 (1848))—the type of which, from Hong Kong, is in the National Collection—
though included by Miss Ricardo among the synonyms of 7. striatus, is in reality
perfectly distinct. Although allied to T. striatus, Fabr., it is, in the ¢ sex, readily
distinguishable, inter alia, by the greater extent of the area of enlarged facets in the
eyes, and by the absence of all trace of a dark band on this area.

The description of T. striatus given by Wiedemann (Auss. Zweifl. Ins., 1, p. 155
(1828)) would appear to apply better to 7. tenens, Walk., than to the true T. striatus,

Fabr.
9. Tabanus virgulatus, sp. n. (fig. 5).

Q—Length (onespecimen) 16mm. ; width of head,5-75 mm. ; width of front at
vertex, 0-6 mm. ; length of wing, 13-5 mm.

Fig. 5. Head of Tabanus virgulatus, Austen, sp. n., 2 ; a, front view; 6, profile.

Medium-sized species, with dark olive-grey thorax relieved by lighter stripes and
dorsum of abdomen blackish-brown, bearing a sharply defined, light buff, longitudinal
median stripe, and on each side of this, between it and lateral margin, a longitudinal
series of somewhat fainter, light buff ovoid blotches or spots, diminishing successively
in size towards posterior extremity, and disappearing before actually reaching tt.

Head : Front and subcallus olive-buff pollinose, former clothed with minute blackish
hairs above, and with similar pale hairs below ; face, jowls and occiput pale smoke-
grey pollinose, clothed with whitish hair, hind margin of upper part of posterior orbits

+ Fabricius (Joc. cit.) in his description of T. striatus writes: ‘‘ Abdomen fuscum lineis tribus
albis, lateralibus, a basi ad medium ductis, media a medio versus apicem.”’

SOME SIAMESE TABANIDAE. 447

fringed with very short and inconspicuous yellowish hair; front in 2 of moderate
width, which diminishes very slightly from above downwards, length of front equal
to about six times its breadth between lower inner angles of eyes, vertex exhibiting a
median, blackish, elongate mark, but ocellar tubercle wanting ; a single frontal callus
(see fig. 5), which is blackish-brown, bottle-shaped in outline and sharply defined,
relatively fairly broad yet clearly though narrowly separated from each eye, its
lower margin straight and on a level with lower inner angles of eyes, its upper extremity
produced into a stout, tapering extension reaching to a point a little above half the
length of the front ; eyes bare (no trace of a band or bands visible in dried condition,
in case of type) ; palpi pale pinkish-buff, proximal segment clothed with whitish hair
(its outer surface—except tip, perhaps tinged with neutral grey), distal segment
considerably swollen at base, then tapering to a slender point, its outer surface clothed
with minute, appressed whitish or yellowish-white hairs, mixed with minute, appressed
black hairs ; antennae cinnamon-rufous, annulate portion of third segment (at least
in case of type) blackish-brown, distal half of expanded portion infuscated on outer
side, first segment light greyish pollinose above, clothed with minute black hairs on
upper surface and with yellowish hairs below, its upper distal angle considerably
produced and embracing second segment, latter small, its distal extremity ringed with
minute black hairs, its upper distal angle small and sharp, not projecting beyond that
of first segment, expanded portion of third segment in 9 narrow and elongate (see
fig. 5), with a sharp, prominent angle on upper margin a little beyond end of proximal
third, length of annulate portion equal to about three-fifths of that of expanded
portion. Thorax : Dorsum of scutum longitudinally striped with light olive-grey, in
manner frequently seen in genus Tabanus—a short median stripe on front border,
not extending quite half-way to transverse suture, and on each side of this, between
it and lateral margin, a somewhat broader, uninterrupted stripe, reaching to posterior
margin of scutum ; lateral borders of scutum pale olive-grey, dorsal surface of scutel-
lum unrelieved by lighter markings, its lateral margins light drab ; dorsum, including
scutellum, clothed with a mixture of minute, appressed cream-buff or yellowish hairs,
and short, fine, erect blackish hairs; swelling occupying depression at each end of
transverse suture mouse-grey, and clothed mainly with black or blackish hair; a
strip on each side, commencing above base of wing and including lower border of
post-alar callus, clothed with whitish or silvery-white hair ; pleurae and pectus pale
smoke-grey, clothed with whitish hair. Abdomen: Median stripe on dorsum commen-
cing on first (visible) segment and terminating abruptly on hind margin of sixth,
diminishing somewhat in width posteriorly and on second to fourth tergites, inclusive,
clearly constituted in each case by a narrow truncate triangle, with its base resting
on hind margin ; ground colour of stripe smoke-grey pollinose ; stripe thus formed is
clothed with minute, appressed, glistening Naples yellow hairs, the combined effect
being light buff ; second to fifthtergites, inclusive, each with a light buff, longitudinally
ovoid spot on each side, midway between median stripe and lateral margin, these
spots or blotches similar in composition to the median stripe itself, but those on fifth
segment very small and faint ; sixth tergite on hind margin on each side with a small
patch of appressed Naples yellow hairs, representing and in continuation of the spot
on the preceding segment, more distinct vestiges of actual spots being visible on sixth
segment when abdomen is viewed at a low angle from behind ; ground colour of sides
of second to sixth tergites, inclusive, cinnamon-rufous ; lateral extremities and hind
margin of seventh tergite cream-buff, clothed partly with glistening Naples yellow
hair, and partly with longer black hair; lateral extremities of first (visible) tergite
pallid neutral grey pollinose (posterior angles faintly pinkish-buff) and clothed with
whitish hair, lateral extremities of following five tergites whitish pollinose, clothed
with glistening whitish or silvery white hair, which towards and on hind margins of
posterior segments may merge into glistening Naples yellow hair ; dorsum, except as
stated, clothed with minute, appressed black hairs; venter, except last segment,
cinnamon-drab or light cinnamon-drab, with a more or less distinct though not sharply
defined median, quadrate, mummy-brown blotch (resting on front margin but not

448 MAJOR E. E. AUSTEN.

reaching hind border) on each segment from second to sixth, inclusive ; first (visible)
ventral scute with a smaller, blackish, median quadrate blotch, which can be seen
between the hind coxae ; seventh sternite mouse-grey, with a darker median blotch,
smaller than that on preceding segment ; hind margins of second to sixth ventral
scutes, inclusive, more or less distinctly cream-buff ; entire venter with a thin pollinose
covering of pale smoke-grey, which, when abdomen is viewed at a very low angle
from behind, entirely conceals the median blotches ; median blotches on fifth and sixth
ventral scutes clothed with semi-erect black hair ; terminal segment, except towards
lateral extremities of hind margin, clothed as usual with coarse, erect black hair ;
venter, except as stated, clothed with short, appressed, glistening whitish or yellowish
hair. Wings : Tinged with mouse-grey ; veins chiefly mummy-brown, in places, such
as base of third and proximal portion of fifth longitudinal, paler ; stigma elongate,
almost colourless and scarcely distinguishable. Squamae smoke-grey or drab-grey,
with brownish borders clothed with pale hair. Halteres: Knobs cream-coloured,
stalks sepia-coloured or paler. Legs: Coxae pale smoke-grey, clothed with whitish
hair ; femora, except extreme tips, black or olivaceous black, with a pale, smoke-grey,
pollinose covering, which is, however, largely wanting on the inner side of those of
the front legs, and in case of middle and hind femora is denser on lower portion of
outer surface than elsewhere ; extreme tips of front femora cream-buff, those of middle
and hind pairs cinnamon-coloured, all femora clothed with whitish hair; front
tibiae, except distal third, which is blackish brown, cream-buff, clothed with minute,
appressed, glistening cream-coloured hairs mixed with minute black hairs; middle
and hind tibiae pinkish-buff (their extreme tips reddish brown), middle tibiae clothed
similarly to those of front pair with a mixture of minute cream-coloured hairs and
black hairs, and with longer black hairs on posterior margin of extensor surface ;
hind tibiae clothed mainly with minute, appressed, glistening cream-coloured hair,
and having on outer margin of extensor surface a fringe of medium length composed
partly of black, partly of cream-coloured hair; front tarsi black, middle and hind
tarsi dark brown or blackish-brown above, proximal segment in case of middle and
hind tarsi somewhat paler at base; all tarsi clothed above with minute black hairs,
second and following (particularly third and fourth) segments of front tarsi consider-
ably expanded, second and following segments of middle tarsi also somewhat expanded.

BANGKOK, November, 1919 (F. J. Godfrev).

The species just described, which is allied to Tabanus striatus, Fabr., and to
T. tenens, Walk. (see above, p. 445), is distinguished from the former of these, in the
© sex, by the shape of the frontal callus ; by the median longitudinal stripe on the
dorsum of the abdomen being as fully developed on the second segment as on the
following ones ; by the outline of the third and fourth segments of the front tarsus,
viewed from above being rounder ; and by the distal portion of the expanded part
of the third antennal segment being narrower and more elongate. From 7. tenens,
Walk., the new species is distinguishable in the 9 sex owing to the shape and darker
coloration of the frontal callus, and to the expanded portion of the third segment
of the antennae being more attenuate.

Tabanus virgulatus is also very closely akin to T. rubidus, Wied., and to the
Indian T. priscus, Walk. (syn. T. albimedius, Walk. .). AS regards the former, it
would seem that, in addition to the differences given in the Key above (see p. 435),
IT. virgulatus is distinguishable in the 2 sex owing to the interval between the lower
portion of the frontal callus and the eye on each side being distinctly narrower.
It is quite possible that, when further material in good condition and from different
countries is available for study and comparison, it will ultimately be found that both
T. virgulatus, Austen, and T. priscus, Walk., are forms of a single widely distributed
species, namely 7. rubidus, Wied.

10. Tabanus rubidus, Wied.
Tabanus rubidus, Wiedemann, Diptera Exotica, i, p. 69 (1821).

SOME SIAMESE TABANIDAE. 449

Two 99, foot of Doi Sutep (alt. circa 1,200 ft.), near Chiengmai, 7,9,iv.1921.
Dr. Barnes’s field-note on these specimens runs: “Caught while attacking my
horse ; a very common fly about horses in this locality.”’

As shown by the series of specimens of this species already in the British Museum
(Natural History), the range of Tabanus rubidus extends at any rate from Bombay
to Hong Kong, and includes Nepal, Burma, the Federated Malay States, Singapore,
Cochin China and Annam. In Singapore, according to a note by Mr. P. S. Falshaw,
T. rubidus bites cattle as well as borses ; a 9 from Pahang, Federated Malay States,
2,500 ft., 1916 (7. R. Hubback—presented by the Imperial Bureau of Entomology),
was taken on a specimen of Rhinoceros sumatrensis, Cuv.

11. Tabanus pugnax, sp. n. (fig. 6).

2.—Length (two specimens) 13-4 to 15-2 mm. ; width of head, 4-5 to 5 mm, ;
width of front at vertex, just over 0-5 mm.; length of wing, 12-75 to 13-4 mm.

Eyes bare, apparently with three purple bands; a single elongate frontal callus
in 9; dorsum of thorax, including scutellum, olive pollinose overlaid with greyish,

Fig. 6. Head of Tabanus pugnax, Austen, sp. n., Q ; a, front view ; 8, profile.

and uniformly clothed with appressed, glistening Naples yellow hair mixed with fine ,
erect black hair; dorsum of abdomen, brownish-tawny or brownish ochraceous-tawny,
with a more or less conspicuous, median, longitudinal stripe extending from front margin
of second (visible) to hind margin of fifth tergite, and composed of a continuous serves
of truncate, ochreous pollinose triangles, clothed with appressed, glistening buff-vellow
hair; fourth and following tergites, otherwise than as already stated, and with exception
of hind margins and posterior angles, mainly blackish-brown.

Head: ¥ront ochreous pollinose, clothed, at least on upper half, with minute,
fine black or blackish hairs, subcallus, face, jowls and occiput somewhat greyer
(dark olive-buff pollinose), sides of face clothed with fine blackish hair, occiput,
jowls and remainder of face clothed with pale yellowish hair, hind margin of occiput
fringed above with very short and inconspicuous ochreous hairs, interrupted behind
vertex by a series of somewhat longer fine black hairs ; front in 2 somewhat narrow
or of medium breadth, slightly broader above than below, its length equal to about
74 or 8 times its breadth between lower inner angles of eyes, a narrow, dusky, indistinct,
elongate triangular mark on vertex, but no trace of an ocellar tubercle ; frontal
callus (see fig. 6 a), black, bottle-shaped, its base on a level with lower inner angles of

450 MAJOR E. E. AUSTEN.

eyes, its upper extremity produced into a lanceolate prolongation extending to a
point about two-thirds of length of front above lower margin of latter, bottle-shaped
portion marked with a narrow, vertical, median groove ; proximal segment of palpr
egreyish-olive pollinose, clothed with fine, yellowish hair, distal extremity of proximal
segment vinaceous-buff, sometimes with a few black hairs on outer side above, terminal
segment pale cinnamon-brown, elongate and bluntly acuminate, but little swollen
at base, clothed on outer surface with minute, appressed black hairs, and with a
few minute yellowish hairs on proximal half of under surface; first and second
segments of antennae cinnamon-drab, clothed with minute black hairs, first segment
also with some yellowish hairs below, this segment considerably swollen distally and
partially embracing second segment, upper distal angle of latter noticeably produced,
third segment ferruginous or orange-cinnamon (terminal annulus blackish-brown in
case of type), its expanded portion in @ fairly deep at base and not particularly
elongate {in case of type, about one-third longer than annulate portion), with a
blunt, but well-developed proximal angle on upper margin. Thorax: Dorsum
somewhat more greyish on anterior border, immediately behind head ; swelling in
depression at each end of transverse suture agreeing in coloration with remainder
of dorsum, but clothed for most part with black or blackish hair, upper and outer
surfaces of post-alar calli also clothed mainly with black hair; pleurae and pectus
egreyish-olive pollinose, clothed with yellowish hair. Abdomen: Hind margins of
third to fifth tergites, inclusive, narrowly ochreous pollinose, and clothed, at least in
part, with short appressed hairs of same colour as those on median stripe ;_ posterior
angles of same three tergites and lateral extremities of sixth tergite ochraceous-
tawny or cinnamon-buff ; hind borders of sixth and seventh tergites more or less
distinctly ochreous pollinose ; second and third tergites sometimes with a more or
less distinct blackish-brown blotch (not reaching hind margin, and larger in case of
third tergite) on each side of median stripe, third tergite also with a faint blackish-
brown blotch, not reaching lateral margin, at each lateral extremity ; hind margin
of first (visible) tergite, in median line, with a tiny patch of appressed, glistening,
buff-yellow hair ; lateral borders of first five tergites fringed with fine, pale buff-yellow
hair; hind margin of sixth tergite with a few glistening buff-yellow hairs in median
line ; dorsum, except as stated, clothed with minute, appressed black hairs ; venter
cinnamon-coloured, extreme base smoke-grey, remainder when viewed at a low angle
from behind seen to be suffused with a dark olive-buff pollinose covering, which is
especially pronounced on last four segments ; hind margins of second and following
sternites deep olive-buff, and clothed with minute, appressed, glistening buff-yellow
hairs (sometimes wanting in case of last segment) ; ground colour of last two sternites,
except hind margins (and sometimes extreme lateral extremities of one or both),
blackish-brown ; second (or third) to fifth sternites, inclusive, sometimes each with a
more or less distinct, iron-grey or dark olive-grey, quadrate median blotch, resting
on base of segment in each case, but not reaching hind margin, in other cases ground-
colour of fourth and fifth sternites, except lateral extremities and hind borders,
mainly dark olive-grey ; venter, except last segment and apart from what has
already been stated, clothed partly with black, partly with appressed, glistening
buff-yellow hair, the latter perhaps predominating on second and towards lateral
extremities and hind borders of the four following sternites ; terminal sternite as
usual clothed for most part with longer, coarser, and more erect black hair. Wangs:
Strongly tinged with sepia ; veins mummy-brown ; sf/gma tawny-olive, narrow and
elongate or sometimes inconspicuous. Sguamae sepia-coloured, with darker
borders fringed with pale hair. Halteres mummy-brown, stalks and tips of knobs
sometimes paler. Legs: Coxae, greyish-olive, trochanters brownish-grey, clothed in
each case with fine, pale yellowish hair ; femora, except extreme tips, blackish-brown
or olivaceous-black, more or less densely covered, at least in case of upper and outer
sides of middle and hind femora, with greyish-olive pollen, and clothed with hair
similar to that on coxae, tips of femora cinnamon-buff ; tibiae cinnamon-coloured

SOME SIAMESE TABANIDAE. 451

(front pair sometimes cinnamon-buff), their distal extremities dark brown, front and
middle tibiae clothed mainly with buff-yellow hair, distal extremities in both cases,
and distal two-thirds of extensor surface of middle tibiae clothed with black hair,
flexor surface of hind tibiae, except at tip, clothed with bright ochreous or ochraceous-
orange hair, extensor surface of hind tibiae fringed with black, or black mixed with
ochreous hair, tips of hind tibiae clothed with black hair; tarsi blackish-brown
(first segment of middle tarsi more or less reddish brown), clothed above with minute
black hairs, third and fourth segments of front tarsi in female moderately expanded.

Doi Chom Chang, near Chiengmai, alt. 5,500 ft., 12.iv.1921. Of the holotype
and the single paratype of T. pugnax, Dr. Barnes writes :—‘‘ These flies attacked
me at about 10 o'clock a.m., at an altitude of 5,500 ft. ; this species is not uncommon.”’

What would appear to be a variety of 7. pugnax is represented in the National
Collection by two 2° from Siam (precise locality unknown), taken at light, 3.111.1914
(K. G. Gairdner). These specimens, which are in poor condition, shrunken and
partly denuded, having originally been preserved in spirit, differ from the typical
form as described above mainly in the coloration of the femora, which are largely
or chiefly cinnamon or pinkish cinnamon-coloured, although, in the case of one
specimen, those of the hind legs are in places strongly tinged with brownish.

The species just described is allied to Tabanus fulvimedius, Ric. (Rec. Ind. Mus.,
Calcutta, iv, p. 197 (1911)—nec Walk.), of Formosa, and to T. fulvimediordes, Shirak,
(Blood- Sucking Ins. Formosa, Pt. 1, Tabanidae, Taihoku, p. 219, pl. v, fig. 8, pl. x1
ngs. 6, 7 (1918)), which is stated by its author to be fairly common in the south of
Japan. From both of these, however, in the 2 sex at any rate, T. pugnax is distin-
guishable by its narrower front ; by the frontal callus (what is the lower callus in
T. fulvimedioides, Shir., in which there are two frontal calli) being narrow and elongate,
with its upper extremity—instead of suddenly and abruptly contracted, as in
T. fulvimedius, Ric.—smoothly and directly continuous with the raised, lanceolate
ridge representing the upper callus; and by the expanded portion of the third
segment of the antenna being deeper and shorter.

12. Tabanus pugiunculus, sp. n. (fig. 7).

§.—Length (one specimen), 13 mm. ; width of head, 4:5 mm. ; length of wing,
10-6 mm.

Fig. 7. Head of Tabanus pugiunculus, Austen, sp. n., §; a, front view ; 6, profile

Dorsum of thorax, including scutellum, deep greyish-olive pollinose, uniformly clothed
with minute, appressed, glistening Naples ° yellow hairs, mixed with fine, erect, cream-
coloured hair; dorsum of abdomen russet, fifth to seventh segments, inclusive, except
lateral and hind borders, olivaceous black, a simoke- grey pollinose, and from certain angles

452 MAJOR E. E. AUSTEN.

not very conspicuous median longitudinal stripe, clothed with minute, appressed, glistening
Naples yellow hairs, extending from base of second (visible) to hind margin of sixth
tergite ; wings hyaline, with conspicuous cinnamon-brown stigma.

Head: father less than upper half of frontal triangle in g¢ mummy-brown,
remainder of frontal triangle pale olive-buff pollinose, face, jowls and occiput pallid
neutral-grey pollinose, clothed with whitish hair, hind margin of upper border of
occiput in g without any noticeable fringe of hair ; outer extremity of area of enlarged
facets in eye of 9 broadly and bluntly rounded off (see fig. 7), enlarged facets them-
selves conspicuously coarse, and below and at outer extremity of area sharply
differentiated from smaller facets ; posteriorly transition to smaller facets is more
gradual, though a border, fairly uniform in width, of small facets runs up to and
reaches vertical triangle ; palpz, vinaceous-buff (proximal. segment somewhat infus-
cated on outer side), clothed with hair like that on jowls, interspersed with which
on outer side of distal segment are scattered, minute black hairs, distal segment
cylindrical, bluntly pointed at tip, not conspicuously swollen; first segment of
antennae in 3 light ochraceous-buff, somewhat swollen distally but not embracing
second segment, clothed above with minute black hairs and on outside and below
with pale yellowish hairs, second segment ochraceous-tawny, its distal margin clothed
on outer side with minute black hairs and its upper distal angle but little produced
in g, third segment tawny (in case of type, distal extremity of expanded portion
dark brown and last two annuli missing), expanded portion in ¢ elongate and some-
what narrow, with blunt but prominent angle on upper margin near base. Thorax :
Dorsum showing indistinct traces of three longitudinal and incomplete, impressed,
dusky lines ; swelling in depression at each end of transverse suture pale ochraceous-
buff pollinose, clothed with fine cream-coloured hair; swelling on each side above
base of wing, behind end of transverse suture, pallid mouse-grey pollinose ; pleurae
and pectus, at least in type, blackish mouse-grey, clothed with fine, whitish hair.
Abdomen: Lateral extremities, except lateral borders, of second to fourth tergites
inclusive, at least in case of type, more or less dark brown ; lateral and hind borders
of fifth to seventh tergites, inclusive, as well as posterior angles of fourth tergite,
cinnamon-buff or ochraceous-buff ; lateral borders of first six tergites with a more
or less distinct, pallid neutral-grey pollinose covering, and clothed with pale, whitish
hair ; first (visible) tergite, except narrow median area immediately behind scutellum,
which is clothed with fine Naples yellow hair, and except its lateral borders, clothed
with minute, appressed black hairs; second to sixth tergites, inclusive, except
median stripe as described in diagnosis above, and except lateral borders, clothed
with minute, appressed black hairs, interspersed with minute, appressed glistening
Naples yellow hairs; seventh tergite in g clothed exclusively with black hairs ;
3d genital appendages clothed above with black hair, sides of eighth segment in 3
clothed partly with whitish or yellowish, partly with black hair ; first three (visible)
ventral scutes and base of fourth orange-cinnamon-coloured, last three ventral
scutes and rather more than posterior half of fourth ventral scute, except hind
border or hind margins in case of fourth to sixth scutes inclusive, olivaceous black ;
first three ventral scutes each with an ill-defined dark mouse-grey or olivaceous-
black median blotch, not reaching hind border in either case, but otherwise quadrate
and extending full length of segment in case of first two visible segments, transverse
and occupying less than posterior half in case of third segment ; posterior borders
or margins of third to sixth ventral scutes inclusive cream-buff ; entire venter light
neutral grey pollinose, second (visible) and three following segments, as well as
hind margin of sixth segment clothed with appressed cream-buff hair, last two
ventral scutes clothed with erect black hair, coarser as usual in case of last scute,
and mixed with minute, appressed cream-buff hairs in case of penultimate scute.
Wings : Costa blackish-brown, veins otherwise mummy-brown. Squamae light drab,
borders mummy-brown. Halteres, knobs blackish-brown or clove-brown, stalks
tawny-olive. Legs: Front coxae light neutral grey pollinose, middle and hind coxae _

SOME SIAMESE TABANIDAE, 453

neutral grey or deep neutral grey pollinose, all three pairs of coxae clothed with
whitish hair ; front femora olivaceous-black, neutral grey pollinose on outer side
below, clothed with fine black hair, which on lower border of outer side of distal
half is mixed with whitish or yellowish white hair ; middle and hind femora mouse-
grey (their distal extremities greyish cinnamon-coloured), clothed, except outer
surface of middle femora, with whitish hair, outer surface of middle femora clothed
with black hair; tibiae, except rather less than distal half in case of front pair,
and rather less than distal fourth in that of middle and hind pair, ochraceous-tawny,
distal extremities of front tibiae blackish-brown or olivaceous-black, those of middle
and hind tibiae reddish-brown, tibiae clothed with black hair (flexor surfaces in case
of middle and hind pairs clothed with appressed cinnamon-buff hair), extensor
surfaces of hind tibiae (at least in 3) fringed with fine and fairly long black hair ;
tarsi black (middle pair missing in case of type), first segment of hind pair paler
(sepia-coloured), all tarsi clothed with minute black hairs.

Near Bangkok, vi.1921, caught in a railway carriage. With reference to this
species, Dr. Barnes writes :—‘‘ I have seen a number of specimens of this fly.”’

What is quite possibly the opposite sex of T. pugiunculus to that of the type
is represented in the National Collection by a solitary 9 taken at Phrapatoon, in
August, 1906 (Dr. P. G. Woelley). While agreeing with the type in many respects,
such as in the presence of a pale (Naples yellow-haired) longitudinal median stripe
on the dorsum of the abdomen, in the character of the wings (including the shape
and coloration of the stigma), and in having strongly infuscated femora, this speci-
men exhibits certain differences—notably in the expanded portion of the third seg-
ment of the antennae being much deeper and entirely tawny (not dark brown at the
tip); in the dorsum of the abdomen being mainly dusky (fuscous, or olivaceous-
black), instead of russet with a dusky distal extremity ; in the venter (apart from
hind margins of segments) being uniformly neutral grey, without any trace of dark
median blotches ; and in the pale portion of the front tibiae being clothed with
glistening yellowish (Naples yellow) hair. The front is relatively rather broad
(4 to 44 times as long as its breadth between the lower inner angles of the eyes) ;
and there is a single, rather large, mummy-brown /rontal callus, in shape resembling
an isosceles triangle, with its lower margin just above the level of the lower inner
angles of the eyes, and its upper extremity produced into a narrower extension,
nearly one-third of the front in width, which dies away at a point about one-third
of the length of the front from the hind margin of the vertex. The dimensions of
this specimen are as follows : length, 10-5 mm.; width of head, 3-6 mm.; width
of front at vertex, 0-5 mm. - length of wing, 9 mm.

As regards species lable to be confused with 7. pugiunculus, it may be mentioned
that the British Museum (Natural History) contains three gg of a (possibly undes-
cribed) species of Tabanus from Lower Burma (base of Dawna Hills, 4.111.1908,
Dr. N. Annandale), which at first sight closely resemble that described above. These
specimens, however, may be distinguished by iter alia, their smaller heads, the
absence of a definite area of greatly enlarged facets in the eyes, and their pale
femora.

Finally it may be remarked that Tabanus pugtunculus presents a certain super-
ficial resemblance to the Javanese J. cinerascens, Big., but is distinguishable, inter
alia, by its infuscated femora.

13. Tabanus agnoscibilis, sp. n

2,—Length (one specimen) 10:4 mm. ; width of head, 3-2 mm. ; width of front
at vertex, 0-4 mm.; length of wing, 9 mm.

Small species, with apparently bare eyes, narrow front and single frontal callus in &,
dorsum of thorax uniform deep greyish-olive, and tapering abdomen which, except last

454 MAJOR E. E. AUSTEN.

three segments, which are blackish-brown, is ochraceous-tawny above, dorsum of ,
abdomen also with a paler, median, longitudinal stripe.

Head: Subcallus and front in 2 except vertex, tawny-olive pollinose, face, jowls,
occiput and vertex pale smoke- -grey pollinose, upper extremities of sides of face
adjoining subcallus faintly tinged with tawny-olive ;- jowls and hinder edge of posterior
orbits below clothed with pale yellowish hair, fringe of similar hair on hinder edge of
upper part of posterior orbits so short as to be scarcely discernible ; front in 9 only
very slightly narrower at lower than at upper extremity, and without trace of an
ocellar tubercle; frontal cailus tawny-olive, elongate and roughly longitudinally
cylindrical-ovate in outline, with its lower end just above lower inner angles of eyes,
and its upper extremity produced into a linear extension, the reddish-brown ter-
mination of which reaches to a point about half-way up the front ; palpz pinkish buff,
clothed on outer side with minute, appressed black hairs, proximal segment below
with longer yellowish hair, distal segment in 9 elongate acuminate, moderately swollen
at base; first and second segments of antennae cinnamon-coloured (first segment
somewhat pallid neutral grey pollinos e above), clothed on outer surface with minute
black hairs, first segment expanded as usual from base to tip, but its upper distal
extremity not enveloping second segment, upper distal angle of latter not conspicu-
ously produced (third segment wanting in case of type). Thorax: Dorsum, including
scutellum, clothed with short, glistening, appressed Naples yellow hair, lateral
border of dorsum and dorsal surface of scutellum smoke- -grey pollinose, swelling
occupying depression at each end of transverse suture with a vinaceous-buft ground
colour, and sparsely « clothed below with longer, erect black hair; pleurae and pectus
smoke-grey pollinose, and clothed with fine whitish hair. Abdomen: Paler, median,
longitudinal stripe light pinkish-cinnamon pollinose on ochraceous-tawny, smoke-grey
pollinose on blackish- brown portion, extending from hind margin of first (visible) to
tin margin of sixth tergite, inclusive, and clothed with minute, appressed, glistening

Naples yellow hairs ; lateral borders of first six tergites somewhat smoke-grey pollinose,
and clothed with hair similar to that on median stripe ; dorsum otherwise than as
already stated clothed with minute, appressed black hairs ; first four ventral scutes
pinkish-cinnamon, clothed with minute, appressed, elistening Naples yellow hairs,
last three ventral scutes blackish-brown : - fifth and sixth ventral scutes somewhat
greyish pollinose, clothed with appressed blackish hair, mixed on fifth scute with
minute, glistening Naples yellow hairs, seventh ventral scute clothed as usual with
coarse, erect black hair. Wangs: Hyaline, with cinnamon-brown or paler veins ;
stigma ochreous, elongate, not conspicuous. Squamae light drab or drab-grey, with
pale-haired borders. Halteres cream-buff, stalks somewhat darker. Legs: Coxae
smoke-grey or mouse-grey pollinose, clothed with whitish hair; front legs, except
coxae, blackish brown, distal fourth of front femora and rather less than. proximal
half of front tibiae cinnamon-coloured ; middle and hind legs, except coxae, ochraceous-
tawny, bases of femora and upper surface of tarsi, except practically whole of first
and extreme base of three following segments, more or less dark brown ; front legs,
apart from coxae, mainly clothed with black hair, posterior border of upper surface
of femora clothed with fine, yellowish hair; middle and hind legs clothed mainly
with vellowish (yellowish Ww hite) hair, upper ‘surface of hind femora and outer edges
of extensor surfaces of hind tibiae also with minute black hairs; front tarsi not
expanded.

SIAM (precise locality uncertain), 3.ii.1914, “at light” (K. G. Gairdner).

The type of this species might easily be regarded as a 9 of T. pugiunculus, Austen,
were it not for the exceedingly pale stigma, differently coloured halteres and much
less infuscated femora. Should it after all prove to be conspecific with the 3 that has
been treated as the type of 7. pugiunculus, the 2 provisionally regarded as belonging
to that species will represent a new one.

SOME SIAMESE TABANIDAE. 455

14. Tabanus siamensis, lic.
Tabanus siamensis, Ricardo, Rec. Ind. Mus., iv, p. 212 (1911).

This species, the provenance of which is given by its author simply as “ Siam,”
is not represented in the series obtained by Dr. Barnes. The type and paratype,
both of which are completely denuded as wel! as somewhat shrivelled, and were
evidently preserved in spirit before being pinned, are in the National Collection.
A moment's glance at either specituen is sufficient to show that what Miss Ricardo
describes as the “shining Llack”’ colour of the thorax is simply due to excessive
denudation-—doubtless caused by washing about while in alcohol. The pollinose
covering of the dorsum of the thorax, of which vestiges are still conspicuous, is
isabella-coloured, so that, although not the slightest trace of hair is now discernible
on the dorsum, it is clear that, whatever be the actual colour of the latter in this
species, it is certainly not “ shining black.” :

457

NEW SPECIES OF AFRICAN SIMULIIDAE AND FURTHER STUDIES
On THE EARLY, STAGES:

By 2. WwW. |) Pomeroy, M.B.., F.EsS.,
Government Entomologist, Nigeria.

(Plates XIII & XIV.)

It has been stated by Malloch and other authors that the superficial differences
between the various species of Simulium in the adult stage are often so minute as
to be discernible only after a very careful study of the genus. Previous studies
had led the writer to believe that the characters of the pupal respiratory filaments
would prove to be of constant specific value, but further investigation has caused
this view to be modified with regard to the actual branching, though the structure
of the chitinous wall and the general appearance still seem to be constant specific
characters.

In the case of one species described in this paper, S. hirsutum, and the varieties
S. hirsutum var. dubium and S. hirsutum var. adersi, though the branching of the
filaments varies very considerably, even in the same pair of filaments from the same
individual pupa, yet the chitinous structure of the filament wall appears to be the
same in all specimens.

The difference in the position and actual number of the branches seems quite
understandable from a morphological point of view and may be explained on the
supposition that the filaments primarily arise from three main stems, which form
into a single stem at the base. The pupa of the var. dubium, though bearing the
same number of branches as the type that has been selected as the true Airsutum,
namely eight, differs in that the branching takes place, in the case of the second
and third main stems, at a very considerable distance from the base.

These two forms were taken from the same locality and were in the greatest
number amongst the specimens obtained, though individuals were taken at the time
showing considerable variation in the distance of the branching point from the base.
With regard to the form adersi, this was obtained from a very different locality and
altitude, but it will be seen from the figure given (Pl. xiv, fig. 6) that the first
main stem has developed a further branch, the second main stem divides again into
two more branches and in the case of the third main stem the branching appears
to have retracted until the three branches arise from nearly the same point. The
position of the branching of this form appears to vary very considerably in a series
of specimens, but no difference can be observed in the chitinous structure of the
outer wall. The male genitalia, wing venation, hind claws of the female, and the

general appearance and colour of the adults of all these three forms appear to
be the same.

A similar case of variation in the pupal branching occurs in the species S. alcocht,
S. alcocki var. violaceum and S. alcocki var. coalitwm, described in this paper. The
adults of these varieties appear to differ slightly in colour, but in no definite
structural character.

The actual position of the branching of the filaments may depend a great deal
on the growth of the larva. Taylor* states that the rudimentary pupal filaments are
developed in the very young larvae at the same time as the imaginal rudiments ;
on this point the writer is fully agreed.

* Taylor, T. H., Trans. Ent. Soc., London, 1902, pp. 701-716.

458 A. W. J. POMEROY.

Two pupae of entirely different species have been obtained by the writer in which
the pupal filaments coalesce for a very considerable distance before branching,
forming a peculiar long main stem. In the case of one of these pupae, described in
this paper as S. alcocki var. coalitum, the imago was well developed within, and the
male genitalia were dissected out, proving identical with those of the true S. alcock?.

As regards the other aborted form, the imago was not sufficiently developed to
allow of a positive determination, but from the number of the branches and the
chitinous structure of the outer wall, it seems to be a form of S. hirsutum. It is
rather remarkable that only single specimens of such aborted pupae were found
among some hundreds of pupae collected and examined, and that the same type of
malformation occurred in two very different species. It is suggested that this
may be due to some adverse condition during the growth of the larval stage.

The writer is now engaged on a monograph of the genus, and from present
knowledge it would seem that the inter-relation of the species may be best determined
from the combined characters of the wing venation, the male genitalia, the hind
claws of the females and the chitinous structure and general appearance of the
pupal filaments.

Simulium hirsutum, sp. nov.

3. Length, 1-4 mm. Antennae black, covered with short grey pubescénce.
Thorax velvet-black, covered with deep golden pubescence. Pleuvae brown, lacking
patch of soft hairs on membranous area. Wangs hyaline, radius unforked. Abdomen
velvet-black, with iridescent patches on the 2nd, 5th and 6th segments, abdominal
scale bearing a fringe of long yellow and brown hairs, the first three segments covered
with golden pubescence dorsally. Legs: front legs rich brown, the femora covered
with dull golden hair ; hind legs, coxae brown, with a few pale yellow hairs, remainder
of leg rich purple-brown, covered with brown hairs and a few scattered golden hairs,
second tarsal joint with deep excision near base. Gevtalia (PI. xiii, fig. 1) : basal pieces
large ; claspers about half the length of the basal pieces, very peculiarly constructed,
the interior margin being extended and folded over into a rectangular flap, the apex
of the clasper pointed and bearing a “ bill-shaped ” finger-like process ; anal plates
not very well defined and bearing a fringe of stout bristles on the outer margin ;
adminiculum very broad, bearing a well defined pouch covered with short curved
hairs arising from well defined pits; arms ending mesally in a single rather short
and very stout spine turned outwardly.

Habitat.—Described from six specimens reared from pupae found attached to
grass blades in a swift-flowing mountain stream.

TANGANYIKA TERRITORY : Morogoro, 21.x1.1917.
Type in the British Museum.

©. Length, 1-5mm. Head: frons and face dark purple-grey, covered with golden
pubescence ; antennae dark purple-brown, covered with dark grey pubescence.
Thorax and scutellum very dark purple-brown, almost black, covered with golden
pubescence. Pleurae brown, lacking patch of soft hairs. Wangs hyaline, radius
unforked. Abdomen deep brown, thickly covered with light golden hairs. Legs:
front legs deep purple-brown, femora and tibiae covered with coppery pubescence ,
hind legs purple-brown, the pubescence darker at joints and interspersed with a few
light yellow hairs, especially on the tibiae and metatarsi; tarsi dark brown, second
tarsal joint with excision near base, claws with a prominent tooth at base.

Habitat—Bred from pupae from same locality and on same date as male type.
Not found biting.

Pupa, type form (Pl. xiv, fig. 4).—The respiratory filaments are eight-branched,
arising from three main stems : the first dividing dichotomously ; the second dividing

—

NEW SPECIES OF AFRICAN SIMULIIDAE. 459

dichotomously and then dividing once again, forming three in all; and the third
dividing dichotomously and once again in the same way but nearer the base. This
is a very common type.

Pupa var. dubium, nov. (PI. xiv, fig. 3)—The filaments are eight in number,
but the second stem divides again very much more distally from the base, and the third
main stem divides again at about two-thirds the entire length from the base.

The typical pupa and this variety were taken from the same locality and on the
same date as the male type.

Pupa var. adersi nov. (Pl. xiv, fig. 6)—The respiratory filaments are 11 in
number, arising from three main stems at base : the first divides into three branches ;

the second divides into three, the first two again dividing dichotomously, making

five in all; and the third divides into three very near the base. The position of the
branching of the third main stem varies greatly, even in the filaments on either side
of the same individual. The same number and general arrangement seems very
constant in this form, no really intermediate forms having been obtained as yet.

East AFRICA: Zanzibar, 22.vii.1917, from grass blades in small stream.

The genitalia of the males, the female structure and the general coloration of
the adults appears to be the same in all specimens bred from or dissected from the
three types of pupae.

Simulium on sp. nov.

3g. Length, 1-5 mm. Antennae dark brown, covered with grey pubescence, the
first two te ae naked and distinctly reddish-brown. Thorax and scutellum,
very dark velvet-brown, covered with light golden pubescence. Pleurae fuscous,
lacking patch of soft hair. Wungs hyaline, radius unforked. Abdomen velvet-black,
basal scale bearing a long fringe “of pale yellow hairs, a diagonal lustrous blue stripe
on either side of the 2nd, 5th and 6th segments. Legs: ficnit legs, coxae and femora
straw-coloured, dark brown at apex, tibiae straw- coloured, dark “br own just at basal
joint, more so at apex, tarsi dark brown; hind legs, coxae yellow, femora yellow,
brown at apex, tibiae yellow, dark brown at base and apex and along outer margin,
metatarsi pale yellow, dark brown at apex and along inner margin, remainder of
tarsi brown, second tarsal joint with slight excision near base. Genitalia (Pl. xii, fig. 4):
basal pieces large; claspers about two-thirds the length of the basal pieces, rather
thick, tapering at apex, which bears a short finger-like process ; anal plates long,
rather cup-shaped at apex and bearing numerous hairs and bristles ; adminiculum
broad, apical margin forming a lip covered with a fringe of short curved hairs ; arms
ending mesally in a single long strong spine, turned back outwardly.

Habitat——Bred from pupae attached to grass blades in slow-moving stream.
NIGERIA: Ibadan, 500 ft., 6.x1.20.

Type in the British Museum.

Q. Length,1-6mm. Head: frons and face grey, covered with silver pubescence ;
antennae fuscous, covered with short grey pubescence, the first two and part of the
third segments deep orange. Thorax dark lustrous grey, completely covered with
dense brassy pubescence in the case of freshly emerged specimens, silvery towards
outer margin. Pleurae brown, lacking patch of soft hairs. Wings hy aline, radius
unforked. Abdomen dark brown, the first six segments covered with golden hair,
the 7th and 8th tergites rather bare, shining, with a few sparse black bristles. Legs :
front legs, coxae yellow, femora and tibiae yellow, dark brown at joints, tarsi brown ;
hind legs, coxae yellow, femora yellow, black at apical joint, tibiae brown, yellow
at base and banded yellow across middle, metatarsi pale yellow, dark brown at apex,
remainder of tarsi black, second tarsal joint with excision near base ; claws with
prominent tooth at base. >

(5296) 2L

460 A. W. J. POMEROY.

Habitat.—Bred from pupae found at same locality and date as male type. Not
found biting. :

Paratype in the British Museum.

Pupa, type form (Pl. xiv, fig. 7)—Respiratory filaments seven-branched, arising
from three main stems. The first main branch divides dichotomously at some distance
from base ; the second divides dichotomously and once again, making three branches
in all; the third divides dichotomously at some distance from the base, but arises
in conjunction with the second rather than from the main base. .

Pupa var. vielaceum, nov. (PI. xiv, fig. 8).—The respiratory filaments differ
from those of true S. alcocki in being ten-branched, and arise from the main base
at a wider angle. The third main stem subdivides into five branches in all.

The adults appear to differ slightly in coloration, the males having a more pro-
nounced violet area on the sides and on the dorsal portion of the first segment of

the abdomen. There appears to be no difference in the characters of the male
genitalia.

Hatbitat—Pupae obtained and adults reared from same locality and on same
date as the type of S. alcocki, also on subsequent dates.

Types in the British Museum.

Pupa var. coalitum, nov. (PI. xiv, fig. 1)—The respiratory filaments of this form
show a very peculiar development. The branching takes place at a very considerable
distance from the main base, there being 10 branches in all. The general arrange-
ment is similar to that in S. alcocki var. violaceum, but the branches seem to have been
welded together as the result of malformation. The genitalia of the male imago,
which was well developed within the specimen obtained, showed no difference in
character from those of the true S. alcockit. More material may show that this
form is a reversion or modification of some earlier type from which both the forms
S. alcockt and S. violaceum have evolved, especially as a similar case occurs with
S. hirsutum.

Simulium divergens, sp. nov.

3. Length, 1-7 mm. Antennae light brown, covered with fine light pubescence-
Thorax deep velvet-brown, covered with light golden pubescence. Pleurae light
brown, lacking patch of soft hair. Wings hyaline, radius unforked. Abdomen
deep velvet-brown, almost black, covered with light golden hairs dorsally, lower
surface of abdomen light brown. Legs: front legs with coxae and femora honey-
yellow, tibiae light brown at base and apex, tarsi very dark brown, almost black ;
hind legs with coxae yellow, femora yellow, with very dark brown area at apex,
tibiae dark brown, pale at basal joint and across middle, tarsi almost black, second
tarsal joint with excision near base. Genitalia (PI. xiii, fig. 3): basal pieces broad ;
claspers about two-thirds the length of basal pieces, the distal portion turned almost
at right angles, the apex of the outer margin pointed and bearing a very short single
finger-like process ; anal plates broad, inner margin with a fringe of short spines ;
adminiculum very broad, bearing a narrow pouch in centre covered with short hairs
arising from distinct pits; arms very strong and ending mesally in a single very
strong blunt spine, turned outwardly, behind which lies a thin membranous area
more strongly defined along the dorsal margin.

Habitat—Bred from pupae attached to grass blades in slow-moving stream.

NIGERIA: Ibadan, 4.xii.1920, 500 ft.

I'vpe in the British Museum.

Described from a single specimen bred from pupa and from a specimen dissected
from pupa.

NEW SPECIES OF AFRICAN SIMULIIDAE. 461

Length, 1-7 mm. Head with frons and face dark brown, pollinose, covered
with pale yellow pubescence; antennae dark brown, covered with minute
pubescence, Ist, 2nd, and part of 3rd segments honey-yellow. Thorax black, with
dull greenish tints, covered with light brassy pubescence, silvery towards sides,
the usual lyre-shaped vittae prominent; scutellum covered with very long pale
brassy hairs. Pleurae dark brown, lacking patch of soft hairs. Wangs hyaline,
radius unforked. Abdomen dull brown, first three segments covered with dull golden
hairs dorsally, last three segments shining dorsally anid covered with sparse yellowish
hairs. Legs: front legs with coxae yellow, femora and tibiae yellow, dark brown
at apical joints, tarsi brown, almost black ; hind legs with coxae yellow, femora
yellow, but dark brown at apex ; tibiae dark brown, pale yellow at apical joint and
diagonally across middle ; basal two-thirds of metatarsi yellow, remainder of tarsi
brown, almost black, second tarsal joint with excision near base, claws with prominent
tooth at base.

Habitat and locality the same as for male type. Not found biting. Described
from a single specimen bred from an isolated pupa.

This species is closely allied to S. aureostmile, Pomeroy, but differs insome of the
characters of the male genitalia, the coloration of the female, and in the structure
of the pupal filaments.

Pupa.—Cocoon strong, but somewhat loosely woven, of the wall-pocket pee
but without “side openings.’’ Respiratory filaments four-branched (PI. xiv, fig. 2).
The first two arise from a main stem, which is narrowed at base. The remaining
pair branch from a stem at a short distance from the main base. The angle between
the filaments is very wide. The ends of the branches are rounded and_ the
chitinous wall is rather weak. The surface of the chitin, which is covered with
minute nodules, is very different in appearance from that of S. auveosimile. Pupa
described from the specimen from which the male type emerged.

Simuiium vorax, sp. nov.

Q. Length, 2-5mm. Head with frons and face grey, covered with shining yellow-
grey pubescence ; antennae dark brown, covered with very short fine grey pubescence,
the Ist, 2nd and part of 3rd segments clear brown-orange. Thorax dark lustrous
grey, brown-grey at sides, the lyre-shaped vittae very dark, prominent and curved ;
the entire surface in fresh specimens covered with thick light greenish- golden
pubescence ; scutellum covered with long greenish-golden hairs. “Pleuvae brown- erey,
pollinose in some lights, lacking patch of Sett hairs. Wungs hyaline, radius unforked.
Abdomen dull woken black, thickly covered with shining yellow-grey pubescence
arranged in whorls, the last three segments Jess dull, and fhe 8th and 9th segments
with long pale yellow-grey hairs interspersed among the pubescence. Legs: front
legs, dark brown, the coxae and basal half of the femora and basal two- ahr of
tibiae covered with shining yellow-grey hair, tarsi dark brown, almost black ; hind
legs dark brown, basal half of femora and basal two-thirds of tibiae covered with
shining yellow-grey hair, tarsi rich dark brown, almost black, basal half of metatarsus,
with exception ‘of inner margin, covered with shining yellow-grey hair, second tarsal
joint with excision near base, claws simple, with no tooth,

Habitat—Taken biting voraciously and engorging on donkey near stream.
Described from 35 specimens taken at same time and place; jg unknown.
TANGANYIKA TERRITORY: Amani, 14.x1.1917.

Type in the British Museum.

This species is very near to S. neavet, Roub., but differs especially in size, and
in the shape and colour of the thorax, which is uniformally dull black in the latter

species.
(5296) ag Oa

462 A. W. J. POMEROY.

Simulium unicornutum, Pomeroy.*

6. Length, 1-5 mm. Antennae fuscous, heavily covered with silver-grey
pubescence. Thorax velvet-black, covered with thick golden pubescence ; meta-
thorax slate-blue and scutellum black, both covered with thick golden pubescence
and long golden hairs. Plewrae fuscous, lacking patch of soft hairs. Wangs hyaline,
radius unforked. Abdomen velvet-black, with iridescent violet-blue patches on 2nd,
5th and 6th segments. Legs: front legs with coxae, femora and tibiae honey-yellow,
dark brown at the apex of the tibiae and a dark brown spot at the apex and near
the base of the femora, tarsi black; hind legs with coxae fuscous, femora honey-
yellow, dark brown at apex , tibiae dark brown, banded yellow at apex and across
middle: basal two-thirds of metatarsi pale, remainder of tarsus black, second tarsal
joint with excision near base. Genitalia (Pl. xiii, fig. 2): basal pieces rather broad ;
claspers a little longer than the basal pieces and rather tapered towards the apex,
which bears a single finger-like process ; anal plates rather short, bearing a clump
of bristles at apex and a few stout hairs at base ; adminiculum very broad, concave
at the centre, which bears a patch of numerous small curved hairs ; arms very strong,
ending mesally in a single long spine turned outwardly. }

Habitat—Bred from pupae attached to grass blades in small stream.
NIGERIA: Ibadan, 500 ft., 6.viii.1920, 9.xii.1920, 11.x11.1920.

Described from specimens bred from isolated pupae corresponding in all details
to the type. Material placed in the British Museum.

©. Length, 2-6 mm. Head with frons and face silver-grey, with light golden
pubescence ; antennae fuscous, covered with fine grey pubescence, Ist, 2nd and
part of 3rd segments dull orange. Thorax and scutellum very dark grey, covered with
light greenish-golden pubescence. Pleurae dark brown, lacking patch of soft hairs.
Wings hyaline, radius unforked. Abdomen dark brown, basal scale with fringe of
long golden hairs, the three basal segments covered with golden pubescence, the
last three segments shining dorsally. Legs : front legs with coxae and femora honey-
yellow, tibiae yellow, banded dark brown just above base and at apex, tarsi dark
brown ; hind legs with coxae and femora honey-yellow, latter dark brown at apex,
tibiae dark brown, banded yellow at base and across middle; basal two-thirds of
metatarsus yellow, remainder of tarsus dark brown, second tarsal joint with excision
near base, claws with prominent thick tooth at base.

Described from many specimens of same date and from same locality as males
and bred from isolated pupae corresponding in all details to the type. Not
found biting.

Material in the British Museum.

Simulium palmeri, sp. nov.

3. Length, 1-5 mm. Antennae black, covered with short grey pubescence.
Thorax: prothorax velvet-black, covered with thick golden pubescence, increasing
in area laterally and diminishing toward the median line ; mesothorax velvet-black,
covered with thick dark purple-brown iridescent pubescence ; metathorax shiny
slate-blue, dotted with a few sparse black hairs: scutellum fuscous, covered with
dark hairs. Pleurae fuscous, lacking patch of soft hairs. Wangs hyaline, radius
unforked. Abdomen velvet-black, covered with black hairs; an iridescent violet
patch on the sides of the 2nd, 5th and 6th segments. Legs: front legs with coxae
fuscous ; femora fuscous, middle portion heavily covered with silvery pubescence,
brassy in some lights ; tibiae covered with silvery pubescence, dark brown at apex ;
tarsi almost black and very hairy ; hind legs with coxae black ; femora deep purple-
brown: tibiae light golden-yellow at base, remainder deep purple-brown, with a

* Pomeroy, Ann. Mag. Nat. Hist. (9) vi, 1920, p. 79, pl. ili.

NEW SPECIES OF AFRICAN SIMULIIDAE. 463

band of golden-yellow pubescence across basal third; tarsi almost black, second tarsal
joint with excision near base, claws with distinct tooth at base. Ge nitalia very similar
to those of S. unicornutum, Pomeroy ; the genera! size appears to be smaller and the
adminiculum not so long and the styli more prominent.

NIGERIA: Ubiaja, 15.1.1921.

Type in the British Museum.

Described from pupae found in swift hill-stream, alt. 900 ft. Bred from isolated
pupae.

Q. Length, 1-5mm. Head: antennae fuscous, covered with silver- -grey pubescence,
Ist, 2nd and part of 3rd segments honey- yellow ; frons bare, slate-blue pollinose ;
face fuscous-blue pollinose in some lights, bare except for a few black bristles. Thorax
dark brown, shining, covered with “dark brown pubescence and with a few golden
hairs, more numerous in front and laterally ; scutellum fuscous, covered with long
black bristles. Plewvae fuscous, lacking pate h of soft hairs. Wings hyaline, radius
unforked. Abdomen very dark brown, almost black, thickly covered with black hairs,
last four segments shining dorsally. Legs: front legs with coxae fuscous, femora
brown, with a pale vellow area at middle and along frontal margin, tibiae dirty
yellow, banded dark brown just below base and at apex, tarsi dark brown: hind legs
with coxae vellow, femora dark brown, paler at middle, tibiae dark brown, pale yellow
at basal joint and with an oblique yellow band across middle ; basal two-thirds of
metatarsus yellow, remainder of tarsus brown, second tarsal joint with excision near
base, claws with thick prominent tooth.

Described from specimens reared from isolated pupae taken on same date and
from same.locality as male type. Not found biting.

This species differs distinctly from S. wnicornutum, Pomeroy, in the darker colour-
ing of the legs, the colour and pubescence of the thorax, especially t the scutellum,
and also in the colour of the frons and face. There is no appreciable difference
in the male genitalia, except in size and possibly in the adminiculum. The pupa,
however, is very distinct and constant in the formation of the respiratorv filaments,
and on the several different characters of this and the adult stage the writer considers
it to be a distinct species.

Pupa.—The respiratory filaments are somewhat similar to those of S. unicornutum,
and consist of a single bent tube on either side (Pl. xiv, fig. 5). The angle at the
base, however, is very distinct and more acute, and the tube is constricted into a
series of globes. This globular appearance is ve ry noticeable in the natural state,
and in prepared mounts it may be seen that it is the result of the arrangement of
the chitin forming the walls of the tube. The cocoon is of the wall-pocket type,
very strong and without side openings. Described from many specimens similar
to that from which male type emerged. Taken same locality and date. Material
placed in the British Museum.

EXPLANATION OF PLATE XIV.

Respiratory Filaments of Pupae of Simulinm from West Africa.

Fig. 1. Simulium alcocki, sp. n., var. coalitum, n.

»”?

ade

divergens, sp. N.

hirsutum, sp. n., var. dubium, n.
hirsutum, sp. Nn.

palmeri, sp. N.

hirsutum, sp. n., var. adersi, n.
alcocki, sp. n.

alcocki, sp. n., var. violaceum, n.

Buti. ENT. RESEARCH. VoL. XII. Part 4. PLATE. XIII.

Male Genitalia of West African species of Simulium.

Fig. 1. Simulium hirsutum, Pomeroy, sp. n.
23 3 unicornutum, Pomeroy.
3h - divergens, Pomeroy, sp. n.
4

a; aleocki, Pomeroy, sp. n.

H. Voc. XII. Part 4,

Respiratory filaments of pupze of Simulium,

465

SOME NATURAL ENEMIES OF MANGO LEAF-HOPPERS
(IDIOCERUS SPP.) IN INDIA.

By T. V. SUBRAMANIAM,

Assistant Entomologist, Mysore Department of Agriculture.
(Plates XV & XVI.)

To entomologists in India the three species of mango Jassids—IJdiocerus niveo-
sparsus, I. atkinsoni and I. clypealis-~-are fairly well known, both on account of their
wide distribution and also on account of the serious damage they do to the mango
crop. In the course of our studies on these important orchard pests we have within
the past three years come across three natural enemies of the adult hopper and
one of the nymph. These are (1) a Pipunculid fly; (2) a Stylopid—both internal
parasites; (3) an Epipyropid moth—an external parasite of the adults; and
(4) a Dryinid wasp—an external parasite of the nymphs.

The following are some observations so far made regarding the above-mentioned
parasites.

1. The Pipunculid Fly (Pipunculus annulifemuy, Brun., sp. n.).*

This is a small dark-coloured fly with a large head and large globular compound
eyes of a dark reddish colour (PI. xv, fig. 6). The female fly is provided with a very
sharp, fairly long, honey-coloured ovipositor, which is held pressed against the ventral
side of the abdomen. All the three species of hoppers are parasitised by this insect,
but I. atkinsoni is the least attacked.

Egg-laying by the flies has not been observed as yet. Full-grown and young
maggots have been observed on the hoppers during November and December.
The young maggots are of a rich orange colour and have the peculiar habit of hopping,
like fruit-fly maggots, when placed on a smooth surface.

The parasitised hoppers are very sluggish, the abdomen being much bloated
and tense. They seek dark and shady places underneath the leaves of mango or other
trees growing near by, fix their rostrum to the leaf tissue, and remain in that condition.
The full-grown maggot now makes a way out through the dorsal surface of the
abdomen, between two segments. The maggots drop to the ground and pupate under
the soil. The full-grown maggot (Pl. xvi, fig. 2) is pale-coloured and oval in shape,
with the anterior extremity narrowed and blunt. It is 5 mm. long and 3 mm.
across. The spiracle shows as a small funnel-shaped black depression, a little away
from the extreme posterior end, on the dorsal surface. The pupal period lasts for
14 to 19 days in captivity.

The characteristic appearance of a number of hoppers adhering to the lower surface
of the leaves of mango trees in shady places, with the abdomen partly split asunder,
is a sure indication of the work of this parasite (Pl. xvi, fig. 1).

2. The Stylopid Parasite (Pyrilloxenos compactus, Pierce).

Dissection of a number of mango leaf-hoppers to find out the percentage of
parasitisation by the Pipunculid fly led me to the discovery of this parasite. The
female parasites (Pl. xv, figs. 2,3) are very small, dark brown, elongate ovate creatures,

* For the description of this species, see p. 469.

466 T. V. SUBRAMANIAM.

with two very small tubercular projections, one on each side, at the anterior extremity,
and a semicircular slit-like opening in the middle of the cephalothorax ventrally.
The ventral portion of the cephalothorax behind this slit-like opening is slightly
arched, and its terminal portion is elongated, V-shaped and black in colour. The
dorsal part of the cephalothorax rests against the abdomen of the host, and is as
broad as the ventral part, but only half as long. Its posterior extremity ends in
a very uneven serrated edge, the serrations pointing outwards. Both the V-shaped
ventral and the serrated dorsal end of the cephalothorax, as also the six-segmented
abdomen, are buried permanently in the abdomen of the host. Thus only the anterior
portion of the cephalothorax is found projecting from the pleuro-ventral side of the
abdomen of the host, generally towards the anal end.

As the ova develop and the larvae are formed, the ovisac occupies the major
portion of the abdominal cavity of the host (Pl. xv, fig. 4). Over 1,000 larvae
were counted coming from a single ovisac. These larvae are very small, active,
fish-like creatures of a pale yellow colour, with two long bristles on the last abdominal
segment (Pl. xv, fig. 5); they were observed suddenly shooting out into space
through the slit opening on the ventral side of the cephalothorax of the female,
which projects from the abdomen of the hopper. This was noticed in a hopper a
few minutes after its death. The mode of the entry of the larvae into living hoppers
has not yet been observed.

The male puparium is seen as a small globular brown body, very much resembling
one end of the pupa of a Muscid fly, with a cap anteriorly, on the pleuro-dorsal side
of the hopper (PI. xvi, fig. 6), partly covered by the wings of the latter. The major
portion of the remainder of the pupa is buried in the abdomen of the host. Generally
only one individual is found in a hopper, but sometimes two or three males and
females are found in the same insect, especially in 7. atkinsoni, which species 1s
very heavily parasitised. The male puparium is sometimes found on the ventral side
of the abdomen.

Before the emergence of the adult male, the two large compound eyes and the
head are clearly visible through the pupal shell. The male pushes out the cap-like
anterior extremity and escapes, leaving the pupal case projecting from the abdomen
like a cylinder. The hoppers survive only a few days after the emergence of the male.
The adult males are dark, very slender, active creatures with a pair of large wings
(Pl. xv, fig. 1). The antennae are seven-jointed, the terminal five joints each
broadened and foliaceous. The insects are very active in flight, and are able to
crawl slowly on the sides of a glass beaker. They lived only eight to ten hours
after emergence from the pupae in captivity.

I. niveosparsus has never been found stylopised ; 30 per cent. of J. atkinsoni
and 15 per cent. of I. clypealis were found parasitised in one year. Misra mentions
(Pusa Memoirs, v, pt. 2, 1917, p. 124) a Stylopid parasite of the sugar-cane leaf-
hopper. The parasite of the mango leaf-hoppers appears to be quite different.

3. The Epipyropid Moth (i pipyrops fuliginosa, Tams, sp. n.).*

The oviposition of this moth on the hoppers has not yet been observed. The
caterpillar is small and cream-coloured, and is found attached by the anal end to the
side of the last thoracic segment of the hopper underneath the wings, with the head
facing posteriorly (PI. xvi, fig. 4). The full-grown caterpillar is as long as the abdomen
of the hopper (3-3-5 mm. long and 2 mm. thick), and the body is thick and stout
towards the head, which is found very close to the anal end; but in young larvae
the head reaches only to about one-third or one-fourth of the length of the abdomen.
All the three species of hoppers are parasitised.

* For the description of this species see p. 468.

NATURAL ENEMIES OF MANGO LEAF-HOPPERS. 467

It is not definitely known on what these caterpillars feed. To find this out,
a number of hoppers with the parasites in different stages were minutely observed
in captivity. The larger caterpillars were found to irritate (scrape?) the sides
of the abdomen with the mouth-parts, and the young caterpillars treated in the
same way the soft portions between the abdomen and the thorax dorsally. Very
small quantities of some whitish wax-like matter was noticed all about the cater-
pillars, but no damage to the abdomen of the hoppers was evident. JI am not sure
if they fed on this white waxy matter. The hoppers seemed in no way inconvenienced
by the presence of these parasites on them, and they were alive and quite active for
a long time after the parasites had left them for pupation.

Pupation takes place underneath the leaves of the mango or on the stems, in
a bright white tough silken cocoon (PI. xvi, fig. 3). If the parasites are on J. niveo-
sparsus, pupation takes place on the main trunk and branches where the hoppers
rest ; if they are on the other two species, pupation takes place underneath the leaves.
In 1919 the white cocoons of these parasites were found in such large numbers
underneath the leaves and on the trunks of the trees in a garden in Bangalore, that
they were at first mistaken for a bad attack of the common scale-insect, Pulvinaria
psidii ; but the absence of “ sooty mould ” and the shining appearance on the leaves
were very marked. The caterpillars do not act as a control on the hoppers to the
slightest extent.

4. The Dryinid Wasp.

During the mango blossoming season many of the hopper nymphs were found
moving on the flower -stalks and leaves with small, dark, round objects attached
to the pleural side of the posterior end of the third segment of the thorax, which were
found to be the grubs of a Dryinid wasp. _ There is a dark sac-like covering enclosing
the grub, which is cast off along with the moulted skin of the hopper nymphs.

The full-grown grub is of a rich cream colour, very thick at the posterior end
and tapering to a point anteriorly (Pl. xv, fig. 7). Pupation takes place in a thin
silken cocoon. We have not yet been able to rear the adults from the grubs in
captivity, so that the species has not been identified. In no case were the nymphs
found to survive the attack of the parasite. All the three species of hoppers were
attacked.

* * * ok of * ** u*

The following table shows the percentage of parasitism of the hoppers by the
different parasites.

|
I. niveosparsus. I. atkinsont. | I. clypealis.
1919. 1920. 1919. 1920. | 1919. 1920.
= a = — ee et S| = ESS on.
; 5 D : |
Pipunculid fly... he we sali LOY, ) Very few! 5% Extremely; 10% 2%
few
Stylopid parasite. . 3. Fhe bt) iv Nl Nal erie 30 oy, So7 15% 3%
Dryinid wasp... se oe Poi) cove. Jesstthamn| — 1097 Doe ou, OK
1% |
Epipyropid moth Me or Ae 13% Dok Very Very 10% Nil
rare rare
Total a rs 27) Se/, 3% | 45% 5%, 40% 6%

In conclusion J have to state that the above observations were made by me in
the mango gardens in Bangalore ; the study of the life-history and bionomics is
being continued. I am greatly indebted to Dr. Coleman, the Director of Agriculture
in Mysore, for the ready help and great encouragement shown to me.

DESCRIPTION OF A NEW SPECIES OF EPIPYROPS FROM SOUTH INDIA.
By W. H. T. Tams.

Epipyrops fuliginosa, sp. n. (PI. xv, fig. 8).

Male.—General coloration fuscous, with the vertex of the head, pectus, underside
of abdomen, anal region, legs and base of antennal shaft mouse-grey. Fore-wings,
above, covered with mouse-grey scales, amongst which are scattered irregular patches
of fuscous-black scales, these predominating but forming no definite pattern ;
beneath, covered more uniformly with fuscous-black scales, except in the region
between the first anal vein and the inner margin, which is covered with narrow
elongate oval mouse-grey scales. Hind-wings above and beneath covered with
fuscous scales, the inner margin furnished with long fuscous hairs. Cilia of both
wings fuscous-black.

Female.—Coloration as in male, but fore-wings with more uniform covering of
fuscous-black scales above and beneath, and less mouse-grey scaling on the
underside.

Fig.1. Neuration of Epipyrops fuliginosa,
Tams, sp. n.

Expanse of male 10 mm.; length of body 3 mm.; antennae 2 mm. long, with
twelve bipectinations, the longest two-thirds of length of antennae.

Expanse of female 8 mm.; length of body 3 mm.; antennae 1 mm. long, with
eight bipectinations and one unpaired pectination at base of shaft, the longest
about a quarter of length of antennae.

Fore-wing neuration (fig. 1). The male type and two fragmentary specimens
have veins 7 and 8 connate, and not stalked as in the drawing.

Hind-wing neuration. The costal region has two veins (?6 and 8) apparently
free from base to termen (compare with Perkins’ figure of neuration of Palaeopsyche).
Vein 8 is well developed, ?6 and 5 are very weak. Anal region with the second
anal vein (1b) well developed, the first (la) and third (lc) very weak.

Male type and female, from Bangalore (T. V. Subyamaniam), in poor condition.
Also two males and a female in bad condition.

469

References.

Dyar, H. G. and Strranp, E. Lepidopterorum Catalogus, Part 16, Epipyropidae,
ge. (1913):

FLETCHER, T. B. Proceedings of the Third Entomological Meeting held at Pusa,
3rd to 15th Feb., 1919, p. 979.—Indian Epipyropidae.

PERKINS, R. C. L. Report of the work of the Experiment Station of the Hawaiian
Sugar Planters’ Association, Division of Entomology, Bulletin No. 1, Pt. 2.
1905.—Leaf-hoppers and their Natural Enemies. (Part II. Epipyropidae).

(Colours from: RrimpGway, R. Color Standards and Color Nomenclature.
Washington, D.C. 1912).

A NEW (-PIPUNCULID PAKASITIC’ ON “LEAF-HOPPERS IN INDIA:
By E. BRUNETTI.

Pipunculus annulifemur, sp. n. (Pl. xv, fig. 6).

Head.—-Eyes dark red ; vertical triangle black, frons dull black, lower part with
a somewhat quadrate greyish dust spot ; face dull black, lower half appearing grey
when viewed from above. Antennae dull yellowish, third joint distinctly pointed,
moderately long and almost sub-triangular. Occiput moderately produced behind
eye margins, black, with a little grey dust. Thorax moderately shining black, middle
of dorsum with yellowish brown dust and a little pale pubescence ; pleurae blackish,
with some greyish dust. Abdomen shining black ; a very little yellowish pubescence
towards sides: genitalia with a yellowish tinge. Legs mainly yellow ; coxae black,
also a broad median band on femora (possibly incomplete in some individuals) ;
tarsi darker towards tips ; all femora with a row of very small bristles on about
apical half of underside. Wangs clear; stigma very pale yellow; anterior cross-
vein at about two-fifths of the discal cell; halteres yellowish. Length, 2-5-3-5 mm.

MysorE STATE: Bangalore, about 3,000 ft. (7. V. Subramaniam).

Described from two males in the British Museum.

EXPLANATION OF PLATE XV.

Parasites of Mango Leaf-hoppers (Idiocerus spp.).

Pyvrilloxenos compactus, Pierce, 3.

ventral view.

a) +)

>, dorsal view.

” 3 =:

Different stages of the larva of P. compactus.
Pipunculus annulifemuy, Brunetti, sp. n., &.
Full-grown larva of Dryinid parasite.

Epipyrops fuliginosa, Tams, sp. n., 6.

BUELL. EN. RESEARCH. Vol. “ll Part 4, PLATE XV.

Ba)

ii
WeeeesS
INRESS

Parasites of the Mango Leaf-hoppers.

EXPLANATION OF PLATE XVI.

Parasites of Mango Leaf-hoppers (Idtocerus spp.).

. Lower surface of mango leaf, showing dead hoppers after the

Pipunculus larvae have emerged from them.

. Full-grown larva of Pipunculus annulifemur.

. Mango leaf with cocoons of Epipvrops fuliginosa.

. Adult hopper with larva of Epipyrops attached to its side.

. Abdomen of a hopper, showing the Stylopid parasite im situ with the

triungulins inside it. (The specimen was boiled in caustic potash.)

Abdomen of a hopper with the male puparia of the Stylopid projecting
from it.

. Hopper nymphs, showing larvae of the Dryinid parasites attached to

the thorax.

Butt. ENT. RESEARCH. VoL. XII. PART 4. PLATE XVI,

Parasites of the Mango Leaf-hoppers.

473

SOME NEW INJURIOUS PHYTOPHAGA FROM AFRICA.

By G. E. Bryant,
Entomological Assistant, Imperial Bureau of Entomology.
Family CRIOCERIDAE,

Crioceris viridissima, sp. n. (fig. 1).

Subcylindrical, brilliant metallic green to coppery green, with legs and three
last ventral segments fulvous, and a large fulvous spot on vertex of head.

Length, 6 mm. -

Head about as broad as prothorax, brilliant metallic green, with scattered
punctures and a large fulvous patch on vertex, sulcate between the eyes on apical
half of fulvous patch. Antennae stout, with first four joints with a slight metallic
tinge, the seven apical joints dull black. Prothorax brilliant metallic green, slightly
longer than broad, sides slightly rounded, strongly punctate, with a fovea at middle
near base. Scutellum triangular, green. FElytra brilliant metallic green, punctate-
striate, slightly shagreened, more than twice as long as broad, parallel-sided and
rounded at apex, broader than base of prothorax. Legs fulvous, anterior pair in
the 3 with the tibiae more bent inwards than in 9. Underside with sternum metallic
green, rugosely punctured ; ventral segments of abdomen with the first two metallic
green, apical segments fulvous, sometimes showing traces of metallic green, and
apical margin of second fulvous, with rather long scattered pubescence on all the
segments.

KENyA Cotony: Nakuru, 4.xi.1918, 7 specimens (S. Colclough) ; Migori Valley
S. Kavirondo, 4,200 ft., v.1911, 1 specimen (S. A. Neave). :

Specimens were forwarded by Mr. T. J. Anderson, Chief of Division of Entomology,
Kenya Colony, with the information that they were attacking asparagus.

This new species is a true Cytoceris, approaching in structure more nearly to
European forms such as C. 14-punctata, Scop., than any African species at present
described. It is probably most nearly allied to C. nigropunctata, Lacord., from
South Africa, amongst the African species.

Family HALtIcIDAE.

Cercyonia citri, sp. n. (fig. 2).

Elliptical, convex, black or bluish-black, nitid ; head and thorax finely punctured ;
elytra bluish-black, with a fulvous patch on basal half of each, punctate-striate ;
underside fulvous, legs darker. Length, 4-5 mm.

Head black, finely punctured, with two fulvous spots at base touching anterior
margin of thorax. Antennae inserted wide apart near the inner circumference of
the eyes, reaching just beyond base of thorax, first four joints fulvous, joint 1 equal
to 2 and 3 combined, 5-10 more triangular, with apical half of each darker, apical
joint acuminate. Prothorax black, finely and evenly punctured, more than twice
as broad as Jong, with sides margined (margin in some fulvous) and narrowed towards
apex ; anterior angles produced and acute, posterior margin broadly produced at
the middle. Scwutellum triangular, black (in some reddish). Elytra_bluish-black,
very little broader than base of thorax, about three times as long as thorax, sub-
cylindrical and narrowed posteriorly ; punctate-striate, more feebly towards apex,

474 Guba BRAN

with the intervals finely punctured; a fulvous patch on each extending from
below shoulder to middle, not touching suture or lateral margin. Legs variable,
fulvous to almost black, with femora darker; posterior pair incrassate; a small
spine at apex of hind tibiae. Underside fulvous; sternum strongly punctured ;
first ventral segment more strongly punctured, the rest with scattered punctures
and slightly pubescent.

Male with last ventral segment more sinuate.

Easily distinguished from C. mgricollis, Jac., by its larger size and markings,
which are constant in over 200 specimens before me.

GoLp Coast: Aburi, 8.xi.1916 (W. H. Patterson) ; Ojesu, Ashanti (A. E. Evans).

Reported by Mr. W. H. Patterson, Government Entomologist, Gold Coast, as
a serious pest of all young citrus plants, and widely distributed in the Colony.

Argopistes oleae, sp. n. (fig. 3).

Rounded, convex ; head and prothorax black, finely punctured ; elytra yellow,
with suture and lateral margins black, each with a black vitta ; underside fulvous.
Length, 4-50 mm.

Head nearly hidden in the prothorax, from base to between eyes black, finely
punctured, front and clypeus flavous. Antennae inserted close together, first four
joints flavous, first joint very long and nearly equal to the three following together,

F a
Fig. 1. Crioceris Fig.2. Cercyonia citri, sp. 0.; Fig. 3. Argopistes oleae, sp. n.;
viridissima, sp. N. a, lateral view. a, apex of hind tibia.

last seven joints slightly broader and fuscous. Pvothorax strongly transverse, black,
finely punctured, about three times as broad as long ; sides obliquely converging
and slightly rounded from base to apex, and deflexed ; anterior angles obtuse and
fulvous, posterior margin sinuate. Scutellum black, triangular. Elvira yellow,
with sutural borders and lateral margins black, each with a black vitta extending
almost from base to apex ; finely punctured, longer than broad. Legs: front and
intermediate pairs flavous, hind pair with the femora strongly incrassate, black
with basal part flavous, their tibiae fulvous, broadly dilated, and deeply sulcate,
dentate and terminated by two spurs (fig. 3, a) ; posterior tarsi with first joint longer
than those of the front and middle pairs. Underside fulvous; ventral segments
strongly punctured, with apical segment longer than the two preceding.

CAPE PRoyINcE: 3 99, Cape Town, 9.111.1918.
The larvae are recorded by the Division of Entomology, Pretoria, as mining in
the leaves of olive trees.
This species is a remarkable mimic of the African Coccinellid genera, Dysis and
Alesia.

NEW INJURIOUS PHYTOPHAGA FROM AFRICA. 475

Argopistes sexvittatus, sp. n. (fig. 4).

Rounded, convex, testaceous ; prothorax testaceous, finely and closely punctured ;
elytra fulvous, a little more strongly punctured than prothorax, sutural margin
black, lateral margins broadly testaceous, with inner margin narrowly black from
base of suture to apex, a narrow black vitta down middle of each elytron ; underside
fulvous. Length, 4-40 mm.

The ¢ only differs in its slightly smaller size and ventral segments, the last
ventral being very large, strongly incised and deflexed, and longitudinally sulcate
(fig. 4, a) ; the 2nd, 3rd and 4th segments are much contracted in the middle.

This species is closely allied to A. oleae, sp. n., but differs in having the head
and prothorax testaceous, the elytral punctures are slightly stronger, the lateral
margins are broadly testaceous, with the inner margins black, and the elytral vittae
are narrower and more sinuate.

Fig. 4. Argopistes sexvittatus, sp.n.; a, venter.

Cape PROVINCE: Stellenbosch, 11.x.1920, 2 ¢¢ and 1 2 on wild olive, 3 99 vars.
(Dr. C. K. Brain). Nata: Potgieter’s Farm, i.1900, 1 9 (H. Bell-Marley). ORANGE
FREE STATE: Bloemfontein, 6.11.1916, 2 gg, 4 99 (J. C. Faure).

The species is a leaf-miner on wild olive.

The six specimens from Bloemfontein do not vary iter se, but differ from the
type from Stellenbosch in the black line of the lateral margins extending from the
apex to little more than a third of the lateral margin. Three specimens collected
by Dr. C. K. Brain at Stellenbosch, 11.x.1920, all females, have the elytra and
prothorax blue-black, with a broad testaceous border. These are evidently only
a variety of A. sexvitiatus, as I can find no structural difference.

This species has stood in the British Museum collection since 1867 under the
MS. name of Pseudococcinella sexvittata, Chevr., and I have thought it better to
retain the specific name, as it has probably been widely circulated. This specimen
came from the Hamlet Clark collection, which contained Chevrolat’s collection.

(5296) 2M

477

NOTES ON THE LIFE-HISTORIES OF TWO MESOPOTAMIAN MOTHS.
By Rao Sahib Y. RAMACHANDRA Rao, M.A.,

Assistant Entomologist, Agricultural College, Coimbatore.

Ocnerogyia amanda, Staud., a Pest of Figs.

The following few notes are intended to supplement the excellent account of
the life-history and habits of this insect already published by Mr. P. A. Buxton
(Bull. Ent. Res. xi, pp. 181-186). Working as an Assistant Entomologist under
the Agricultural Directorate at Baghdad, during the year 1919-1920, I had a few
opportunities of observing this pest in Mesopotamia. I met with it in small numbers
at Museyib on the Euphrates in June 1920, and at Karradah (a suburb of Baghdad)
from June to the middle of September of the same year. It is evidently sporadic
in its distribution ; for I did not find it in gardens at Jadriyah, situated only three
miles west of Karradah, nor did I notice it in any of the gardens at Hillah, which
is situated on the Euphrates, about 25 miles south of Museyib.

The eggs are deposited in groups of 20 to 50 or more, being usually laid on the
bark of the stems and less frequently on the underside of leaves. The egg is hemi-
spherical in shape, and is attached by its convex face, while its distal face is flat
and has a central depression. It is shiny yellow when freshly deposited, but soon
becomes covered with dust. The larva when ready to hatch emerges by biting a
large hole in the side of the egg.

The caterpillar when freshly hatched is less than 2 mm. long, light yellow in
colour, and covered with light grey hairs. It passes through six moults before it
becomes full-grown. A caterpillar that hatched on 18.vi.1920 went through the first
moult on 20.vi.20, the second moult on 22.vi.20, the third moult on 24—25.vi.20,
the fourth moult on 27—28.vi.20, the fifth moult on 1.vii.20, and the sixth moult on
4-5.vii.20. It began to build its cocoon on 11.vii.20, and finished it on 12.vu.20,
but unfortunately died on the 13th July. Another caterpillar of the same batch,
which hatched on 18.vi.20, began preparing its cocoon on 11.vii.20, pupated on
12-13.vii.20,and emerged as a moth on 20.vii.20. The larval period in these cases, there-
fore, covered about 24 days. The larva of the last or seventh instar is 22 to 28 mm.
long; it is very hairy. The general coloration of the body is a soft greyish-brown,
marbled with darker markings. The head is rugose and of an opaque pale brown.
The prothoracic shield ig not distinctly defined, but is broad and pale brown with
a median stripe. On each side of the shield there is a prominent, anteriorly directed
tubercle, carrying a conspicuous pencil of spines and hairs. A dorso-median streak
of brownish-orange colour stretches from the head to the hind end. The trunk
carries dorsally a number of tubercles, each bearing a bunch of spinelike bristles and
long silky hairs—all of a greyish colour. The tubercles are disposed more or less
as follows :—(1) A single dorso-lateral row; (2) a double row above the spiracles ;
and (3) another double row below the spiracles.

The larva feeds on the leaves during night-time, and hides in cracks in the soil
or in crevices in the bark during the day. Occasionally, however, a few specimens
may be found resting on the lower surface of leaves during the day. In the
case under my observation at Karradah, most of the leaves were damaged to a
greater or less extent, but the numbers of the pest were not large enough to cause
entire defoliation.

478 RAO SAHIB Y. RAMACHANDRA RAO.

The caterpillar seeks the ground when full-grown and constructs a loose silken
cocoon (into which the hairs of the trunk are incorporated), in cracks of the soil or
of mud walls close by.

The pupa is about 18 mm. long; stout and thickset, translucent grey when
fresh, but turning yellowish-brown after a time. The anterior end is rounded and
stout, while the posterior end is conical and terminates in an elongate spike-like
process, carrying a bunch of recurved hooks at the tip. The wing rudiments reach
almost to the edge of the fourth segment. Dorsally, both on the thorax and on the
abdomen, groups of little chitinous tubercles—each carrying a fairly long curved
grey hair—are noticeable. These form cushions of hair, which serve to support the
pupa as it lies on its back in the cocoon. Similar cushions—but smaller in size
and composed of shorter hairs—are found laterally and ventrally. The moth emerges
in 8 to 11 days in summer, as may be seen from the table given below :—

Pupated. Emerged. No. of days.
(1) 12-13.vii.1920 oe 20-21 .vii.1920 aes 8
(2) 19.vii.1920 sas 30-31 .vii.1920 Bos 11
(3) 22-23.vii.1920 ach 30-31 .vii.1920 sis 8
(4) 23-24.vii.1920 ee Bl vil. vill: 1920).<.. 8
(5) 23-24. vii.1920 me 1—2.viii.1920 ce 9

The duration of the egg stage is not known, but probably varies from a week
to 10 days in summer. Since the larval stage occupied about 24 days in the two
cases under observation and the pupal stage covers 8 to 11 days, the entire life-cycle of
the moth, from the time the egg is laid up to the time of the emergence of the moth,
is about a month and a half. It is therefore probable that there are about three
generations of the pest during the warmer part of the year, 7.e., from April to
September. The insect probably hibernates as a larva in cracks in the soil or in
crevices in the bark: and it is even not unlikely that it passes the winter in the
egg-stage—considering the large size of the eggs. In the neighbourhood of Karradah
the fig caterpillar was not known by any special name except by the general term
“ Dud-et-teen ” (literally, the fig worm).

Banding the stems with tanglefoot might prove effective as well as economical
in large gardens and deserves a trial in affected Jocalities.

Theretra alecto, L.,; on Grape Vines.

A light green shiny spherical egg—1-5 mm. in diameter—was noted on the
10th August 1920, laid on the upper surface of a leaf of a grape-vine, in the neighbour-
hood of the Government tree nurseries at Baghdad. On the 12th August the
egg hatched into a slender greenish caterpillar, 3-4 mm. long, with the anal horn
black and disproportionately long compared with the body. On the 14th August
the first moult took place, and the second on the 17th. The caterpillar was
observed growing rapidly, while the anal horn was noticed to become shorter after
each moult. On the 20th August the third moult was gone through; dots were
noted to have appeared on the sides of the trunk. On the 21st August the cater-
pillar measured about 37 mm., while on the 22nd the length was about 50 mm. On
the 23rd August the fourth skin was cast and the spectacle-like markings were noted
to have appeared. On the 25th August the larva of the fifth or last instar was
about 60 mm. long, reddish-brown, thickset and cylindrical. The anterior part of
the body was tapering, the head being opaque dark brown and small in proportion
to the rest of the body. The segments of the thorax were narrower than those of the
abdomen, but progressively increased in size from the front backwards. Segment 1
of the abdomen was the stoutest, and was marked dorsolaterally with a pair of large
circular yellow-ringed black markings, having the appearance of a pair of eyes.
A similar but smaller and less conspicuous pair of spectacles was noticeable on

NOTES ON THE LIFE-HISTORIES OF TWO MESOPOTAMIAN MOTHS. 479

segment 2. On segments 3, 4, 5 and 6 the spectacles were represented by
elongate, oval, horizontal yellow patches. The sides of the body were yellowish-
brown, with an oblique brown fascia on each segment. The anal horn was 2-5 mm.
long, blunt-tipped, and pinkish-brown in colour. The true legs were pinkish with
white bands, while the prolegs were thick, fleshy and purplish-brown in colour.

On the 28th August the larva ceased to feed and began to construct its cocoon,
which was prepared of leaves and a loose network of a few strong silken strands,
at the bottom of the cage. Under natural conditions the caterpillar presumably
constructs its cocoon in the soil. It pupated during the night of the 30th August.
The pupa was about 2 in. in length, elongate and cylindrical, varying in colour from
a yellowish-brown to a darker brown. Head large ; rudiments of eyes and antennae
clearly impressed : face compressed, prominent and projecting a long way in front
of the eyes. The rudiments of the wings, legs and proboscis extended almost to
the posterior margin of the fourth abdominal segment. Abdomen tapering behind,
terminating in a spike-like chitinous process carrying two strong hooks at the tip;
spiracles large. The moth emerged on the 15th September 1920, being a large insect,
pinkish-brown in colour, with the hind wings and sides of the abdomen rosy red.

The duration of the larval period in the present case was about 19 days, while
the pupal period lasted about 15 days. In a second case, a caterpillar collected in
October 1919 pupated in November 1919, passed the winter in the pupal stage,
and emerged as a moth on the 9th May 1920.

This insect is found on the grape-vine in spring as well as in autumn, but is
usually not seen in large numbers. It is, however, a voracious feeder, a similar
caterpillar kept in a breeding cage having been found to have devoured nearly
100 vine leaves by the time it commenced to build its cocoon.

An allied hawkmoth, Deilephila livornica, is reported by Herr Bredemann to
be a serious pest in vineyards in Upper Mesopotamia (Rev. App. Ent. viii, A,
p. 346).

481

ON THE EGGS AND OVIPOSITION OF PSOROPHORA (JANTHINOSOMA)
LOSTICATA, WiIED> (CULICIDAE):
By Jo Us PAWAN, M.B., Ch.B.,
Trinidad, British West Indies.
(Plate exev LL}

The mosquito, Psorophora posticata, Wied, (Janthinosoma musica, Say), commonly
deposits its eggs in the rain-water that accumulates in the broken cacao pods strewn
in heaps about the cool shady parts of cacao fields.

The ovipositing female assumes a characteristic attitude upon the surface of
the water. The hind pair of legs lie extended backwards to their maximum length
and slightly outwards. The front pair of legs are projected forwards and outwards,
the femur forming almost a right angle with the tibia and the latter an obtuse angle
with the tarsus, which rests forwards and outwards. In both the front and hind legs
the tarsi are the only parts in direct contact with the surface of the water. The
position of the middle pair of legs is very definite ; the femur is directed backwards,
the tibia is acutely flexed forwards and its apex embraces firmly the lateral borders
of the egg-mass, the tarsus projecting sharply backwards and resting upon the sruface
of the water. Whilst the coherency of the egg-mass is maintained by the grip of
the middle tibiae, the buoyancy of the eggs prevents the mosquito from being
submerged during the process of oviposition. This process, which usually takes
place in the morning, Jasts from two to four hours, during which time the insect
seems quite helpless, being unable and unwilling to take wing if disturbed, rendering
herself an easy captive, and giving one the impression of being in pain. After
oviposition has ceased the mosquito continues to rest upon the surface of the water,
often away from the egg-mass, for from two to three hours, then crawls to the side
of the containing receptacle and flies away.

The eggs lie in circular or subquadrate masses, consisting of from 25 to 40 in
number, floating with their long axes perpendicular to the surface of the water
and glued together at their broadest circumference by a gelatinous substance that
helps to keep the mass afloat. A little less than one-third of their length is sub-
merged. The eggs on escaping from the female are of a distinct greyish-brown
colour, but in less than half an hour that portion which is exposed to the air
assumes a dark steel-blue appearance, the whole mass of individual eggs simulating a
honeycomb.

To the naked eye an individual egg (PI. xvii, fig. 1) shows an ovoid shape elongated
at both extremities, with the greatest circumference immediately above the junction
of the blue and brownish portions, the latter tapering rapidly with a sharp and
distinct curve. The lower portion, which retains its colour, is chitinous, while that
which lies above the water is brittle and calcareous.

Under the microscope the portion of the egg that rests above the surface of the
water is seen to be covered with a definite but loosely adherent capsule, studded
with numerous translucent pedunculated and sessile tubercles, arranged in regular
rows and containing air (Pl. xvu, fig. 2). This investing capsule does not extend
beneath the water, and is also absent from a narrow triangular area on the upper two-
thirds of the egg. Along this bare area the brown egg-shell can be seen, for the
change in colour from brownish to dark blue affects the capsule only and not the
egg-shell. Shorn of its capsule the underlying egg-shell is seen to consist of a brownish
outer layer made up of definite circular strands, enclosing an inner thin delicate
layer surrounding the yolk substance, which bathes the nucleus and subsequently
the embryo. No operculum can be seen.

The larva hangs with its head downwards in the floating egg, and in from
eight to ten hours after oviposition ruptures the lower submerged portion and
the longitudinal area of the egg free from investing capsule and then escapes into
the water.

Buti. Ent. REsEancuH, Vou. XIil, Panr 4. PiArE NVA

Fig l. Eggs of Psorophora posticata, Wied.
Fig.2. Egé capsules; note the rows of spikelets.
Fig.3. Eés shells with capsules, after escape of larvae.

483

COLLECTIONS (RECEIVED:

The following collections were received by the Imperial Bureau of Entomology
between Ist October and 3lst December 1921, and the thanks of the Managing
Committee are tendered to the contributors for their kind assistance :—

Dr. W. M. AvERs, Government Economic Biologist :—50 Psochidae; from
Zanzibar.

Mr. T. J. ANDERSON, Chief Entomologist :—73 Siphonaptera, 4 Coleoptera,
11 Mallophaga, 136 Anoplura, 389 Mites, and 15 Ticks ; from Kenya Colony.

Dr. G. ARNOLD :—6 Asilid Flies ; from Rhodesia.
Mr. -E. BALLARD, Government Entomologist :—55 Hymenoptera, 220 Coleoptera,
236 Rhynchota, and 116 Orthoptera ; from South India.

Mr. H. A. Battou, Entomologist, Imperial Department of Agriculture :—
14 Parasitic Hymenoptera, 25 Coleoptera and early stages, 37 species of Coccidae,
1 species of Aleurodidae, and 1 Cimicid bug; from the British West Indies.

Mr. H. W. BEpForD :—64 Coleoptera, 8 Lepidoptera, 5 Crickets, and 1 tube of
Red Spiders on cotton leaves; from the Sudan.

Mr. C. F. C. BEESON, Forest Zoologist :—72 Curculionidae, and 130 Orthoptera ;
from India.

Mr. G. E. Bopkrin, Government Economic Biologist :—132 Culicidae, and

1 Hippoboscid fly; from British Guiana.
Mr. H. E. Box :—60 Chalcids, and 12 Coleoptera and early stages; from
Kenya Colony.
Dr. H. Brauns :—1 Nemestrinid fly and 33 Coleoptera ; from Cape Colony.
Mr. P. A. Buxton :—4 Leptoconops, 6 Tabanidae, 22 other Diptera, 53 Coleoptera,

1 Microlepidopteron, 1 species of Coccidae, 37 other Rhynchota, 160 Orthoptera,
and 1 Chrysopa; from Palestine.

Dr. A. E. CAMERON :—19 Culicidae, 14 Simulium, 46 Tabanidae, 8 other Diptera,
and 5 tubes of early stages ; from Canada.

Mr. L. C. CotEman, Director of Agriculture, Mysore :—10 Diptera and 4 Eppes
cases, 6 Moths, and 12 Rhynchota; from India.

Mr. R. E. Cooper :—A species of gall on Indigofera ; from Burma.

Mr. E. CRESSWELL-GEORGE :—4 Coleoptera, 1 Lepidopterous larva, and 3 Rhyn-
chota; from Nyasaland.

Mr. M. T. Dawe :—2 Tabanidae and 32 Glossina ; from Portuguese West Africa.

Division oF Entomoxocy, Pretoria :—50 Coleoptera; from South Africa.

Mr. C. C. GowbEy, Government Entomologist :—3 Diptera, 25 Parasitic Hymen-

optera, 3 Coleoptera, 5 Lepidoptera, 1 species of Coccidae, 1 species of Aphididae,
and 3 other Rhynchota; from Jamaica.

Mr. H. HarGREAVES, Government Entomologist :—1 Mosquito, 7 Tabanidae,
77 other Diptera, 41 Hymenoptera, 308 Coleoptera, 98 eee 1 Ant-lion,
319 Rhynchota, 70 Orthoptera, and 1 tube of Mites parasitic on a Gryllid ; from
Uganda.

(5296) Qn

484 COLLECTIONS RECEIVED.

Mr. G. F Hi, Entomologist, Australian Institute of Tropical Medicine :—-
2 Hymenoptera and nest, 5 Coleoptera, 2 Lepidoptera, 3 Rhynchota, 6 Orthoptera
and 3 nymphs, 7 Spiders, 1 Centipede, and 1 Millipede ; from Australia.

Capt. Hincston :—10 Crickets ; from United Provinces, India.

Mr. M. Arzat Husain, Government Entomologist :—500 Parasitic Hymenoptera .

and 4 Rhynchota ; from the Punjab.

Mr. R. W. Jack, Chief Entomologist, Department of Agriculture :—2 ‘Poleopteras
from Rhodesia.

Dr. W. B. Jounson :—5 Haematopota, 3 Tabanus, 36 Glossina, and 5 other
Diptera ; from N. Nigeria.

Mr. H. H. Kinc, Government Entomologist :—8 Dipterous larvae and 45
Orthoptera ; from the Sudan.

Mr. N. C. E. MILLER :—85 Hemimeridae ; from Tanganyika Territory.

Prof. S. A. MoxrzEeck! :—5 Coleoptera; from Bulgaria.

Mr. J. C. Moutton :—2 Coccinellidae and 1 pupa, and 31 Orthoptera ; from
Singapore.

Mr. F. Mut
NatTaL MusEum :—403 Orthoptera; from South Africa.
Mr. W. H. Patrerson :—10 Diptera; from the Gold Coast.

Dr. L. Pérrncuey, Director of the South African Museum :—176 Orthoptera ;
from South Africa.

a; from Fiji and Honolulu.

Mr. A. W. J. PomERoy, Government Entomologist :—185 Mosquito larvae,
35 Nycteribiidae, 50. Coleoptera, 71 Lepidoptera, 1 Thysanopteron, 4 Isoptera,
3 Planipennia, 5 Orthoptera, 3 Odonata, 3 Lice, 40 Ticks, and 4 Worms; from
Nigeria.

Mr. A. H. Rircure :—4 Curculionidae ; from the Suez Canal: 1 species of Aleuro-
didae ; from Kenya Colony: and 8 Coleoptera; from Tanganyika Territory.

Mr. H. W. Stumonps :—1 Tabanid, 14 other Diptera, 6 Hymenoptera, 2 Coleoptera,
2 Lepidoptera, and 12 Rhynchota; from Fiji.

Mr. R. VeiItcuH :—51 Diptera, 40 Ants, 54 other Hymenoptera, 257 Coleoptera,
67 Lepidoptera, 5 species of Aphididae, 55 other Rhynchota, 20 Orthoptera, and 12
Neuroptera; from Fiji.

Mr. G. M. VEVERS :—50 Mallophaga; from British Guiana.

Mr. O. H. WALTERS :—46 Coleoptera ; from India.

WELLCOME BUREAU OF SCIENTIFIC RESEARCH :—2 Tabanus, 18 other Diptera,
68 Hymenoptera, 60 Coleoptera, 5 Rhynchota, 12 Orthoptera, and 5 Spiders ; from
various localities:

Mr. G. = Wo tcotTt :—24 Coleoptera ; from Porto Rico.

Mr. R. C. Woop :—159 Culicidae, 5 Glossina, 48 Stomoxys, 66 ieee 123
Simulium, re Psy chodidae, 1 Auchmeromyia, 21 Cordylohta, and 1,275 other Diptera ;
from Nyasaland.

| Title, Contents, Indices to Vol. XI.

BULLETIN OF
ENTOMOLOGICAL
RESEARCH

ISSUED BY THE JMPERIAL
BUREAU OF ENTOMOLOGY.

EDITOR: THE DIRECTOR.

on Ta Cons

Zo MN YOGI!

AS if Ne 8p Y it :
Ae ae dae |

( APRi7: 1923 |

\ a

* f ; Avy

& 4 f y f Oo SP

/HSONI AN US
cane

LONDON :
THE IMPERIAL BUREAU OF ENTOMOLOGY,
41, QUEEN’S GATE, S.W.7.
1921-1922.

485

GENERAL INDEX.

abdominalis, Cephenomyia.
Abies firma, Coccid on, in Japan, 217.
Acacia confusa, Coccid on, in Japan,
212.
Acanthoconops, new subgenus of Lep-
toconops, 3, 24.
albiventris, anatomy of,
7; description of, 26.
x SPInosifrons, Sp. I.,
anatomy of, 5, 6, 7,
9,19; description of,
24-26.
Achaea janata, new Braconid parasite
of in Indias 130; }132:
Achatina fulica, destructive to cotton
in Mauritius, 182.
aconitus, Anopheles.
Acrostichum, Ceroplastes rubens on, in
Seychelles, 127.
actaeon, Hypoderma.
Adalia bipunctata, predacious on Aphids
in Britain, 430.
aderst, Simulium hirsutum.
Adoretus versutus, food-plants of, in
Mauritius, 185, 187, 190.
Aédes, key.to subgenera of, 293; key
to larvae of Palaearctic species
of, 293-295.
+ aestivilis, closely resembling A.
sticticus, 312)
,, (Ochlerotatus) albescens, sp. n.,
in W. Siberia, 309.
fe (Stegomyia) albopictus, breeding-
places of, in Far Eastern
Ports, 402-409; in Palae-
arctic Region, 325.
,, aldrvicht, perhaps a synonym of
An sticiicus, 312.
», (Ochlerotatus) alpinus, in Palae-
arctic Region, 309.
», (Ochlerotatus) annulipes, in
Palaearctic Region, 305;
anatomy of, 298.

(6127)

Aédes argenteus

a”

(Stegomyta fasciata),
not found in Japanese Ports,
402, 408; in Palaearctic
Region, 325 ; breeding in salt
water, 30, 31.

(Ochlerotatus) ashworthi, sp. n.,
in Australia, 75.

aureostriatus, A. japonicus not
considered identical with, 318.

(Ochlerotatus) bancroftianus, sp.
n., In Queensland, 74.

cantans (see A. maculatus).

(Ochlerotatus) caspius, influence
of drought on, in British Isles,
208; in Palaearctic Region,
299 ; anatomy of, 301.

(Ochlerotatus) cataphylla, anato-
my of, 298.

cataphylla var. vostochiensis, in
Palaearctic Region, 310.

(Aédes) cinereus, in Palaearctic
Region, 323 ; in Britain, 427.

(Ochlerotatus) communis, in
Palaearctic Region, 314;
anatomy of, 298.

(Caenocephalus) concolor, 76.

(Stegomyia) cretinus, sp. n., in
Palaearctic Region, 325.

(Ochlerotatus) crucians, 76.

cyprius (see A. lutescens).

(Stegomyia) dendrophila, sp. n.,
in Gold Coast, 74.

(Ochlerotatus) detritus, in Palae-
arctic Region, 310; anatomy
of, 298.

(Ochlerotatus) diantaeus, inPalae-
arctic Region, 311.

(Ochlerotatus) dorsalis, in Palae-
arctic Region, 300 ; anatomy
of, 301.

dorsalis, Theo. (see A. caspius).

(Finlaya) eatoni, in Palaearctic
Region, 319.

486

Aédes (Finlaya)

GENERAL INDEX.

echinus, in Palae-
arctic Region, 320; anatomy
of, 294, 320, 321.

(Ochlerotatus) excyucians, in
Palaearctic Region, 305;
anatomy of, 298.

(Stegomyia) fraseri, 74.

(Ochlerotatus) freyi, sp. n., in
Finland, 298, 301, 306.

(Skusea) funerea, in Amboina,
76.

(Skusea) funerea var. ornata, in
Ceram, 76.

galliv (see Aédes pullatus jugo-
yum).

(Finlaya) geniculatus, breeding-
places of, in Britain, 206-208,
427; in Palaearctic Region,
319; anatomy of, 266, 292, 320.

grahami (see A. dorsalis).

hirsuteron, closely resembling A.
sticticus, 312.

imnuitus (see A. alpinus).

(Ochlerotatus) intrudens, in Palae-
arctic Region, 316; anatomy
of, 301.

(Finlaya) japonicus, breeding-
places of, in Japanese Ports,
405-408 ; in Palaearctic Re-
gion, 318.

(finlaya) koreicus, in Palaearctic
Region, 318.

lazavensis, 309; probably a
variety of A. communis, 315.

(Ochlerotatus) lepidonotus, in
Palaearctic Region, 308;
anatomy of, 292, 301.

(Ochlerotatus) lesne1, perhaps a
synonym of d. sticticus, 312.

(Ochlerotatus) lutescens, in
Palaearctic Region, 306;
anatomy of, 298.

(Ochlerotatus) maculatus (cantans,
waterhousei), in Palaearctic
Region, 304; anatomy of,
298 ; influence of drought on,
in British Isles, 208; breed-
ing in salt water, 30.

(Ochlerotatus) mariae, in Palae-
arctic Region, 303 ; anatomy
of, 294, 301, 302.

metalepticus (see A. pullatus
qugorum).

(Ochlerotatus) nearcticus (see A.
alpinus).

(Ochlevotatus) nemorosus (see A.
punctor var. meigenanus).

(Finlaya) niveus, in Palaearctic
Region, 318.

obscurus (see A. communis).

(Ochlerotatus) palmeni (see A.
communis).

Aédes panayensis, 76.

”

(Ochlerotatus) parvulus, sp. n.,
in Finland, 314; anatomy
Ort, oro).

(Skusea) pembaensis, 76.

pionips, probably a variety of
A. communis, 315.

(Stegomyia) pseudonigeria, 74.

(Ochlerotatus) pulchritarsis, in
Palaearctic Region, 303.

pullatus, anatomy of, 298.

(Ochlerotatus) pullatus var. ju-
gorum, in Palaearctic Region,
316.

(Skusea) punctipes, sp. n., in
Burma, 77.

(Ochlerotatus) punctory, anatomy
of, 298.

(Ochlerotatus) | punctor var.
meigenanus (nemorosus), in
Britain, 208, 427; in Palae-

arctic Region, 313.

(Ochlerotatus) rusticus, in Palae-
arctic Region, 308; venation
of, 285.

(Ochlerotatus) salinellus, in Palae-
arctic Region, 311; anatomy
of, 298.

(Stegomyia) scutellaris (see A.
albopictus).

(Ochlerotatus) semicantans, in
Palaearctic Region, 305;
anatomy of, 298.

(Ochlerotatus) sticticus, in Palae-
arctic Region, 311.

(Stegomyia) sugens (see A. vit-
tatus).

tahoensis, probably a variety of
A. communis, 315.

(Ochlerotatus) tasmaniensis (see
A. crucians).

(Finlaya) togot, breeding-places
of, in Japanese Ports, 405—
408; in Palaearctic Region,
318.

(Ecculex) vexans, in Palaearctic
Region, 322.

vexans var. nipponii, 322.

(Stegomyia) vittatus, in Palae-
arctic Region, 326.

(Ochlerotatus) waterhousei
A. maculatus).

(Ochlerotatus) zammittit,in Palae-
arctic Region, 303.

(see

aegypti, Culex.
aenea, Rachionotomytia.
aestivalis, Aédes.

Africa: w@estrids ‘of 245) 4247-

new

injurious Phytophaga in, 473-475 ;
new Simuliids in, 457-463.

Africa, South, bionomics of Locustana
pardalina in, 155-159.

GENERAL INDEX.

africanus, Phlebotomus minutus.

agnoscibilis, Tabanus.

Agrilus foveicollis, attacking roses in
Bulgaria, 353-354.

Agrilus viridis, 354.

Agromyza phaseoli, on beans in Maur-
itius, 186.

Agyrotis ypsilon, new Braconid parasite
of, in India, 129.

Agyrtes bicolor, a parasite of Scatopse,
231.

aithenit, Anopheles.

alaskaensis, Theobaldia.

albescens, Aédes (Ochlerotatus).

albicans, Culicoides.

albiceps, Chrysomyia.

albigenu, Culex (see C. ventrillont).

albimanus, Anopheles.

albimedius, Tabanus (see T. priscus).

albina, Musca.

albionensis, Orthopodomyia (see O. pul-
chripalpis).

albiventris, Acanthoconops.

albomaculata, Musca.

albopictus, Aédes (Stegomyia).

Alcides crvassus, in the Andamans, 167.

dipterocarpi, sp. n., in seeds
of Dipterocarpus tuberculatus
in India, 166.

Af morto, in South India, 168.
alcocki, Simulium.
aldabraca, Pseudaonidia.
aldrvichi, Aédes.
alecto, Theretra.

Alesia, 474.

Aleurodes vaporariorum
chiton).

Algeria, mosquitos in, 264; Simulium
beckert in, 117.

algeriensis, Anopheles.

Allotheobaldia, subgenus of Theobaldia,
286, 287.

alluaudi, Haematopota.

Alopecurus pratensis, new Aphid on,
in Britain, 430.

alpinus, Aédes (Ochlerotatus).

amanda, Ocnerogyia.

America, Oestrids
in, 249.

America, North, representative mos-
quitos of, 265.

americana, Cuterebra.

americanus, Leptoconops kertészt.

Amblyomma variegatum, on cattle in
Kenya Colony, 235.

amictus, Anopheles.

analis, Cuterebra.

Anastellorrhina auguy (see Calliphora).

Anatrachyntis simplex (see Pyroderces).

Andamans, Alcides crassus in, 167.

andersoni, Culex.

angustifrons, Musca (see M. humulis).

”

(see A stero-

infesting rodents

(6127)

487

angustitarsis, Anopheles.
annandalet, Chionaspis.
annulata, Leucomyia; Theobaldia.
annulatus, Culex.
annulifemur, Ochlerotatus (see Stegomyia
periskeleta) ; Pipunculus.
annulipalpis, Leicesteria.
annulipes, Aédes (Ochlerotatus)
pheles.
annulivostris, Culex ;
Culex taylori).
Anopheles, anatomy of larvae of, 91—
97; Palaearctic species
of, 267-281 : keys to,
269; eggs of, 268, 276.
at aconiius, larva of, in Feder-
ated Malay States, 91,
935596;
ay aitkent, larva of, in Feder-
ated Malay States, 91-97.
Se albimanus, breeding in salt
water, 31.
oe albotaeniatus var. montanus,
larva of, 93.
ies algeriensis, in Palaearctic
Region, 270; egg of, 268.
i amictus, sp. n., in Queens-

; Ano-

Leucomyia (see

land, 71.

ay angustitarsis, 116.

i annulipes, breeding in salt
water, 31; <A. moluc-

censis not a variety of, 71.
ra antennatus (see A. bifur-
catus).
7: argyvopus, possibly a form
of A. hyrcanus, 275.
es astaticus, larva of, 93, 96.
es barianensis, considered dis-
tinct from A. plumbeus,

* barbivostris, larva of, in
Federated Malay States,
91-96.

of bifurcatus, influence of

drought on, in British.
Isles, 205, 206 ; breeding-
places of, in Britain, 427;
in Palaearctic Region, 271;
egg of, 268 ; larva of, 94.

i chaudoyei (see A. multi-
color).

i claviger (see A. maculi-
pennis).

55 cleopatvae (see A. multi-
color).

(Myzomyta) costalis, breed-
ing in brackish water in

Africa, 29> in’ Palae-
arctic Region, 277; egg
of, 268.

crucians, breeding in salt
water, 30.

488

GENERAL INDEX.

Anopheles culicifacies, associated with

”

”?

”

Culex mimulus, 338.
elutus, sp. n., in Palaearctic
Region, 273; egg of, 268.
flaviceps, sp. n., in Egypt,
69

fuliginosus, larva of, 93, 96.

(Myzomyia) hispaniola, 30,
69; in Palaearctic Region,
279 ; egg of, 268.

hyycanus, breeding-places
of, in Far Eastern ports,
402-408; larva of, in
Federated Malay States,
91, 93; in Palaearctic
Region, 274; in Palestine,
116; egg of, 268.

hyycanus var. mesopotamiae,
DIS.

hyvcanus var. pseudopictus,
Pei faye

immaculatus, probably a
form of A. vagus, 70.

karwari, larva of, in Feder-
ated Malay States, 91,
eye e/a

Rocht, larva of, 93, 96.

leucosphyrus, larva of, in
Federated Malay States,
OF.

leucosphyrus var. hackert,
n., in Federated Malay
States, 70.

lewist (see A. maculipennis).

lindesayi, in Palaearctic
Region, 273; breeding-
places of, in Japanese
ports, 405, 406.

ludlowi, breeding in salt
water, 30; larva of, 93.

lukist (see A. algeriensis).

maculatus, 277; larva of,
in Federated Malay
States, 91-96, 97.

maculipalpis, 277.

maculipennis, breeding-
places of, in Britain, 427 ;
influence of drought on, in
British Isles, 205, 206, 208;
in Palaearctic Region,
272; breeding in brackish
water, 29, 31; €g¢" of;
268; larva of, 94; and
malaria, 5; supposed
variety of, in Palestine,
116 (see A. elutus).

malefactoy, breeding in salt
water, 31.

mauritianus, in Palae-
arctic Region, 275.

minimus, associated
Culex mimulus, 338.

with

Anopheles

(Myzomyia) multicolor, in
Egypt and Palestine, 29,
116; in Palaearctic
Region, 280 ; breeding in
salt water, 29, 30; egg
of, 268.

occidentalis (see A. maculi-
pennis).

palestinensis (see A. super-
pictus).

paludis
tianus).

(Myzomyia) pharoensis, in
Palaearctic Region, 277.

pictus (see A. hyrcanus).

plumbeus, breeding-places
of, in Britain, 427; in-
fluence of drought on, in
British Isles, 206—208 ; in
Palaearctic Region, 271 ;
egg of, 268.

pseudopictus (see A.
canus).

pseudopictus flerowi, doubt
as to identity of, 267.

pseudopunctipennis, breed-
ing in salt water, 31.

(Myzomyia) pulcherrimus,in
Palaearctic Region, 277 ;
egg of, 268.

punctibasis, sp. n., 337; in
Japan, 274; breeding-
places of, in Japanese
ports, 405, 406.

punctulatus, characters of,
m0):

punctulatus var.moluccensis,
7:

punctulatus var. tessellatus,
vA

quadrimaculatus, breeding
in salt water, 30.

(Myzomyia) rhodesiensis, in
Palaearctic Region, 278 ;
egg of, 268.

vosst var. indefinitus (see
A. subpictus var. vagus).

sacharovii, doubt as to
identity of, 267.

selengensis (see A. maculi-
pennis).

separatus, 274.

(Myzomyia) sergenti, in
Palaearctic Region, 279.

sinensis (see A. hyrcanus).

(Myzomyia) stephensi, in
Palaearctic Region, 277 ;
breeding in salt water,
30; egg of, 268.

subpictus var. vagus, larva
of, in Federated Malay
States, 91, 93.

(see A. mauri-

hyr-

GENERAL INDEX.

Anopheles (Myzomyia) superpictus, in
Palaearctic Region, 278 ;
in Palestine, 116; Culex
mimeticus associated
with, 338; anatomy of,
278.

superpictus vassilievi, doubt
as to identity of, 267.

- tarsimaculatus, breeding in
salt water, 31.

i tessellatus, larva of, 93, 96.

by. theobaldi, 277.

a turkhudi, 69; breeding in

salt water, 30; egg of,
268.
3 (Myzomyia) turkhudi var.

persicus, n., in Palaearctic
Region, 280.
re umbrosus, larva of, 93.
5 vagus, A.immaculatus prob-
ably a form of, 70.
a willmort, 277.
antancarus, Henicospilus.
Antelopes, Oestrids infesting, in Africa,
246-248.

antennatus, Anopheles (see A. bifur-
catus).

Antestia lineaticollis, bionomics of para-
sites of, in Kenya Colony, 191-201.

antestiae, Hadronotus.

Anthomyia, carrying eggs of Deyma-
tobia hominis, 249; larva of, acci-
dentally producing myiasis, 252.

anthropophaga, Cordylobia.

Antonina bambusae, food-plant of, in

Formosa, 211.
Be. crawit, food-plant of, in
Formosa, 211.

Aonidia obtusa, sp. n., on Verschaf-
feltia splendida in Seychelles, 126.

aonidum, Chrysomphalus.

Aphidius, parasite of Aphis gossypii in
Mauritius, 182.

A phiochaeta ferruginea (see A. xanthina).

5 vujfipes, Causing myiasis,
240, 250; characters of
larva of, 258, 260.
PA xanthina, causing myiasis,
240, 250; characters of
larva of, 258, 260.
Aphis brassicae, on cabbage in Mauri-

tius, 186.
» euonymi, synonym of A.rumicis,
81.

», fabae, synonym of A.rumicis, 81.

» gossypit, on cucurbitaceous
plants in Mauritius, 182, 186 ;
natural enemies of, 182.

re maidis, on maize in Mauritius,
184.

», papaveris, synonym of A. ru-
micis, 81.

489

Aphis rumicis, bionomics of, in Britain,
81-89.
apicalis, Culex ; Cuterebra.
approximata, Cuterebra.
aprepes, Tabanus.
arctica, Theobaldia (see T. alaskaensis).
Areca catechu, Pinnaspis buxi on, in
Seychelles, 127.
argenteus, Aédes (Stegomyia).
Argol, Trigonogenius globulum breeding
IML SSe
Argopistes oleae, sp. n., on olives in
South Africa, 474.
w. sexvittatus, sp. n., on olives
in South Africa, 475.
argyricephala, Lucilia.
Argyroploce rhynchias, on Canavalia
ensiformis in Mauritius, 185.
argyvopus, Anopheles.
Aristolochia acuminata, scale-insects on,
in Mauritius, 182.
Armigeres obturbans, breeding-places of,
in Japanese ports, 405,
406 ; in Palaearctic
Region, 326.
5S ventralis, in Hong Kong, 404.
Artichoke, pests of, in Mauritius, 186.
articulatus, Selenaspidus.
ashworthi, Aédes (Ochlerotatus).
Asia, Eastern, Coccidae in, 211-220.
astaticus, Anopheles.
asininus, Gastrophilus intestinalis.
Asparagus, new beetle attacking, in
East Africa, 473.
Aspidiotus, 218; on Canavalia ensi-
formis in Mauritius, 185.
© inusitatus, Odonaspis peni-
cillata recorded as, 219.
is lataniae, food-plant of, in
Formosa, 219.
A spidomorpha obovata, on sweet potato
in Mauritius, 184.
Asterochiton (Aleurodes) vaporariorum,
colour tropisms of, 355-359.
A theroides, Laingia allied to, 429.
athinsoni, Idiocerus.
atrvox,Cuterebra.
augur, Calliphora (Anastellorrhina).
Aulacaspis cinnamom1, food-plant of, in
Formosa, 212.
- flacourtiae (see Diaspis).
_ pentagona (see Diaspis).
ot vosae, food-plant of, in
Formosa, 213.
Py tegalensis, on sugar-cane in
Formosa, 213.
auvanti1, Chrysomphalus.
aureo-argentatus, Oestrus.
aureosimile, Simulium.
aureostriatus, Aédes.
auribarbis, Cephenomyia.
aurotaenta, Culex (see C. quasigelidus).

490

Australia, bionomics of Euthyrrhinus
meditabundus on mango in, 63-66 ;
new Leptoconops in, 11, 12; house-
flies in, 423-426; mosquitos in,
71, 74,78, 80 ; bionomics of Tabanids
in, 41-62; notes on termites in,
363-399.

australicum, Trichogramma.

australis, Locusta (Pachytylus) (see L.
migratoria ph. danica) ; Musca (see
M. convexifrons).

autumnalis, Musca.

azoviensis, Culex (see C. pipiens).

Bamboo (Bambusa stenostachys),
Coccids on, in Japan, 211, 213, 219.

bambusa, Rachionotomyia.

bambusae, Antonina.

Banana, pests of, in Mauritius, 190.

bancroftianus, Aédes (Ochlerotatus).

Baphinia, Coccids on, in Japan, 2A

barbirostris, Anopheles.

barianensis, Anopheles.

barnesi, Tabanus.

Barraudius, new subgenus of Culex,
328, 332.

basicinctus, Culex.

batchelori, Tabanus (see T. aprepes).

Batocera rubus, on mango in Mauritius,
189.

Beans (Phaseolus vulgare), Astervochiton
vaporariorum on, in Britain, 359;
pests of, in Mauritius, 186 ; Aphis
yumicis on, 81; Phytometra_ ori-
chalcea on, 99.

beckeri, Simulium.

beckit, Lepidosaphes.

beesoni, Sympiezomias.

Bellardia nigrotecta (see Tabanus).

bengalensis, Gastrophilus intestinalis.

bergrothi, Theobaldia (see T. glaphy-
voptera).

bezziana, Chrysomyta (Pycnosoma).

bezzii, Leptoconops ; Musca.

Bibio hortulanus, attacking potatoes in
Germany, 232; larva of, 231.

johannis, bionomics of, in Britain,
224, 231.

,, lacteipennis, bionomics of, in
Britain, 221, 228-230.

marci, bionomics of, in Britain,
221-228.

,, pomonae, larva of, 231.
venosus, bionomics of, in Britain,
221,.229, 230.

bicolor, Agyrtes.

bicyuciatus, Coccus.

bifurcatus, Anopheles.

bimaculata, Uvranotaenia.

bimaculipes, Rachionotomyia.

binotalis, Crocidolomia.

GENERAL INDEX.

bipunctata, Adalia; Forcipomyia.

bitaeniorhynchus, Culex.

bivittata, Musca (see M. humilis).

bjerkandrella, Porpe.

blanchardi, Kirkioestrus.

Blastophaga jacobsoni, bred from figs
in Borneo, 35.

boetica, Lampides.

Bogeria spp., infesting rodents in
America, 249.

Bois d’Amande,
Seychelles, 126.

boisduvali, Diaspis.

bombacis, Rhadinomerus.

Bombax malabaricum, new weevil bred
from, in India, 172.

Boophilus, on cattle in Kenya Colony,
235.

Borneo, new fig insects in, 35—40.

Botys, on Cajanus indicus in Mauritius,

new scale on, in

185.
,, octoguttatus, possibly recorded in
error for Thliptoceras octo-

guttalis on coffee in Mauritius,
187 (and footnote).

bovis, Hypoderma.

brasiliensis, Locusta (Pachytylus) (see
Locusta migratoria ph. danica).

brassicae, Aphis.

braziliensis, Leptoconops.

brevipalpis, Culex ;
Megarhinus.

brevirostris, Osphilia.

Britain, bionomics of Aphis rumicis
in, 81-89; new Aphid in, 429;
Asterochiton vapovarviorum infesting
tomatos in, 355-359; early stages
of Bibionids in, 221-232 ; bionomics
of mosquitos in, 205-209, 427.

broquettii, Grabhamia (see Aédes dor-
salts).

brunnescens, Haematopota.

brunnipennis, Tabanus.

Bubaris indemnis, associated with
termite nests, 371.

buccata, Bogeria.

Bulgaria, Agrilus foveicollis attacking
roses in, 353-354.

Burma, Osphilia egregia in, 180.

Butea frondosa, new weevil bred from,
in India, 178.

buteae, Rhadinopus.

buxi, Pinnaspis.

Cyathomyta ;

Cabbage, pests of, in Mauritius, 186.

cadaverina, Cynomyia.

caesar, Lucilta.

Cajanus indicus, pests of, in Mauritius,
182, 185.

Calandva oryzae, in stored maize in
Mauritius, 184.

GENERAL INDEX.

Calceolaria, Asterochiton vaporariorum
on, in Britain, 359.

calceolariae, Pseudococcus.

calcitvans, Culex ; Stomoxys.

Calliphora, characters of larvae of, 258,

260.

a (Anastellorrhina) augur,
bionomics of, causing
myiasis, 250; larva of,
Dos

5; erythrocephala, causing my-

iasis, 250-252 ; larva of,
254, 255, 257.

ss quadrimaculata, causing my-
lasis, 250.
rf vomitoria, Causing Myiasis,

Z50)= larva ot. 255:
calopus, Culex (see Aédes argenteus).
Calotermes convexus, in Australia, 363.

+ obscurus, in Australia, 363.
camelina, Hippobosca.

Camels, Oestrids infesting, 244, 247.

Canavalia ensiformis, pests of, in
Mauritius, 185.

canicularis, Fannia.

cantans, Culex (Ochlerotatus) (see Aédes
maculatus) .

capensis, Hippobosca ;
Locustana pardalina).

capito, Pachytylus (see Locusta migra-
torta ph. migratorioides).

Capnodium, growth of, due to Coccids,
189.

caprina, Lipoptena.

Carrollia, a subgenus of Culex, 328.

caspius, Aédes (Ochlerotatus).

Cassia fistula, new weevil bred from,
in India, 180.

cataphylla, Aédes.

catoivt, Ceratitis.

Cattle, trypanosomiasis in, in Kenya
Colony, 233-235; warble flies in-
festing, 248.

Cauliflower, pests of, in Mauritius, 186.

cautella, E phestia.

cayennensis, Cutevebra.

centyinifovmis, Rhadinopus.

Cephalomyia maculata (see Cephalopsis
titillator).

Cephalophus spp., Cordylobia vodhaini
causing myiasis in, 243.

Cephalopsis titillator, infesting camels,

Locusta (see

244, 247; characters of larva of,
250:
Cephenomyia abdominalis, in U.S.A.,
250.
iy auribarbis, infesting red
deer, 250.
ss macrotis, in N. America,
Pas Oe
a phobifer, in N. America,
250)

491

Cephenomyia pratti, infesting deer in
N. America, 250.

a stimulator, infesting roe
deer, 250.

re tvompe, infesting reindeer,
250.

* ulvichi, infesting elk, 250.

Ceratitis catoiyi, on citrus and mango
in Mauritius, 189, 190.
Ceratosolen crassitarsus, 38.

a hewittt, sp. n., bred from
figs in Borneo, 35-38.
ap striatus, 38.
Cercyonia citri, sp. n., on citrus in Gold
Coast,-473:

,, nigricollis, 474.

cerviferus, Ceroplastes.

Cerococcus ficoides, food-plant, of, in
Formosa, 212.

Ceroplastes ceriferus, on citrus and

coffee in Kenya Colony,
103.
i: vubens, food-plant of, in
Formosa, 212; on fern
in Seychelles, 127.
cervi, Lipoptena.

Ceylon, Osphilia brevirostris in, 180;
scale-insects in, 127-128.

Chaetococcus, considered a synonym of
Antonina, 211.

Chalcis euthyvrhini, sp. n., parasite of
Euthyrrhinus meditabundus in Queens-
land, 65; description of, 67.

chalcomelaena, Lipopiena.

chalcytes, Phytometra.

chaudoyet, Anopheles (see A. multi-
color).

Chilomenes lunata, predacious on Aphids
in Mauritius, 182, 186.

China, *CoccidS ins 222 l4e JG
mosquitos found in ports of, 401-409.

chinensis, Fiorinia.

Chionaspis, on manioc in Mauritius, 184.

_ annandalet, food-plant of,
in Formosa, 213.

ee city1, on citrus in Mauritius,
188.

om dilaiata, on mango in
Mauritius, 189.

+ eugeniae, doubt as to iden-
tity of, 213.

ne subcorticalis, on Cajanus

indicus in Mauritius,
185; food-plants of, in
Seychelles, 127.
Chlovidea obsoleta (see Heliothis).
Choeroporpa, New World representa-
tive of Culiciomyia, 335.
christophi, Megarhinus.
chrysidiformis, Cobboldia.
Chrysoconops, 291.

492

Chrysomphalus aonidum, food-plant of,
in Formosa, 220.
aurantit, food-plant of,
in Formosa, 219;
bionomics of, in
Kenya Colony, 103—
104; on citrus in
Mauritius, 188.
a jicus, on. citrus , in
Mauritius, 188.

Chrysomyia, characters of larvae of,
254, 255, 258.

albiceps, causing myiasis,
250); Jarvaiot-258),260%

45 bezziana, producing myia-
sis in Africa and India,
239,241) 242) larva
of, 259, 260.

EP dux (see C. megacephala).

3 flaviceps, 242.

if macellaria (see
mya).

5 marginale, causing myiasis,
250.

% megacephala, producing
myiasis in India, 239,
242 ; found in intestines
of man, 252)- larva of,
255 ; synonymy of, 250.

* vufifacies (see C. albiceps).

ts vavipes, Causing myiasis,
250 ; larva of, 258, 260.

Chrysops flavocincta, in Siam, 431.

chrysostoma, Sarcophaga.

cilipes, Haematopota.

cinevascens, Tabanus.

cinereus, Aédes.

cingalaisina, Musca.

cinnamomt, Aulacaspis.

Cinnamomum camphora, Coccid on, in

Formosa, 212.
- ceylanicum, Coccid on,
in Java, 212.
circumscriptus, Culicoides.

cityi, Cercyonia; Chionaspis ; Pseudo-
coccus.

Citrus, new beetle attacking, in Gold
Coast, 474; Coccids on, in Japan,
DT OR 2T ES DAE eS a2? )eemiscale=
insect on, in Kenya Colony, 103 ;
pests of, in Mauritius, 182, 187.

clavki, Leucotermes.

claviger, Anopheles

pennis).
cleopatrae, Anopheles (see A. multicolor).
clypealis, Idiocerus.

coalitum, Simulium alcockt.

Cobboldia chrysidiformis, infesting ele-

phants in Africa, 248.

53 elephantis, infesting
phants in India,
larva of, 255, 261.

Cochlio-

(see A. maculi-

ele-
248 ;

GENERAL INDEX.

Cobboldia loxodontis, infesting ele-
phants in Africa, 248.
oi parumspinosa, infesting ele-

phants in Africa, 248.

Coccidae, from Eastern Asia, 211—220 ;

from the Seychelles, 125-128.

Coccus bicruciatus, food-plant of, in

Formosa, 212.
ES elongaius, tood-plants
Formosa, 212.
,, hesperidum, on citrus and coffee
in Kenya Colony, 103; on
mango in Mauritius, 189.
,, MmMangiferae, on mango in Mauri-
tius, 189.
a viridis, food-plants of, in Mauri-
tius, 187, 188.
Cochliomyia, characters of larvae of,
254, 255, 258.
a macellavia, causing Myia-
sis, 240, 250.
ays viridula, causing myiasis,
250.
Coconut, pests of, in Mauritius, 187.
Codiaeum variegatum, Coccid on, in
Japan, 212.
Coelonia solani, on tobacco in Mauritius,
183.

Coffee, scale-insect on, in Kenya
Colony, 103; parasites of
pests of, in Kenya Colony,
191-201 ; pests of, in Mauri-
tius, 187.

Coffee Bug (see Antestia lineaticollts).

columbae, Haemoproteus.

communis, Aédes (Ochlerotatus).

compactus, Pyrilloxenos.

compositus, Oestrus.

comstocki, Pseudococcus.

concinnus, Culex (see Aédes sticticus).

concolor, Aédes (Caenocephalus) ; Lutzia.

conducens, Musca (see M. humiilts).

conjugens, Gyrostigma.

consputus, Rhadinopus.

convexifrons, Musca
maculata).

convexus, Calotermes.

convolvuli, Herse (Sphinx).

Coptotermes, 363.

Pe lacteus, in Australia, 394.

Coquillettidia, subgenus of Taenio-

vhynchus, 291.
Cordylobia anthropophaga, causing my-
1asisy 24 243%
s vodhaini, causing myiasis,
241, 243.

covinnae, Hypoderma.

covvina, Musca.

Corynoneura, 1.

Cosmophila flava, on cotton in Mauri-

tius, 181.
z xanthindyma (see C. flava).

of, in

(see M. albo-

GENERAL INDEX.

Cosmopolites sordidus, on banana in
Mauritius, 190.

costalis, Anopheles (Myzomyia).

Cotton, pests of, in Mauritius, 181, 182.

crassivostvis, Musca (Philaematomyia) .

crassitarsus, Ceratosolen. e

crassus, Alcides.

Cratopus punctum, on coffee in Mauri-
tius, 187.

crawit, Antonina.

cretinus, Aédes (Stegomyia).

Cricetomys gambianus, Cordylobia
vodhaint causing myiasis in, 243.

crinicauda, Culex.

Crioceris nigrvopunctata, 473.

quatuordecimpunctata, 473.

viridissima, Sp. N., On aspara-
gus in East Africa, 473.

cristata, Gedoelstia.

Crocidolomia binotalis, on cabbage in
Mauritius, 186.

crucians, Aédes (Ochlerotatus) ;
pheles.

Cryptodus gvossipes, in termite nests,
oH hk.

””

”

Ano-

Cryptorrhynchus lapathi, stridulatory
apparatus of, 178.
. mangiferae, on Mango

in Mauritius, 190.
Ctenodactylus guna, 5.
Cucumbers, pests of, in Mauritius, 186.
Culex, key to Palaearctic species of,
328-331 ; characters of larvae
of, (92;
», aegypti, 325, 326.
albigenu (see C. ventrillont).
,, andersont, 78.
,, annulatus, 78, 267.
annulatus var. marocanus
Theobaldia longiareolata).
annulivostris, 78.
apicalis, in Palaearctic Region,
336.
auritaenia (see C. quasigelidus).
,, azorviensis (see C. pipiens).
basicinctus, n. n., proposed for
Leucomyia annulata, 78.
bitaeniorhynchus, in Palaearctic
Region, 337.
(Neoculex) brevipalpis, 79, 336.
calcitrans, perhaps a synonym of
C. pipiens, 345.
calopus (see Aédes argenteus).
cantans (see Aédes maculatus).
concinnus (see Aédes sticticus).
crinicauda,n.n., for Culex parvus,
Taylor, 77.
curriei (see Aédes dorsalis).
domesticus, perhaps a synonym
of C. pipiens, 345.
dorsalis (see Aédes).

(see

oi)

493

Culex fasciatus, perhaps a synonym of

C. pipiens, 345.

,. fatigans, breeding-places of, in
Far Eastern ports, 402, 404,
405, 406, 407; in Palaearctic
Region, 345.

», ficalbi (see Theobaldia fumipen-
n1S).

,, filipes (see Rachisoura).

,, flavivostris, perhaps a synonym
of Theobaldia morsitans, 289.

,, flavovirens, believed to be a
Chironomid, 267.

,, fusculus (see Aédes geniculatus).

», (Neoculex) fuscus, 79.

,, hayashi, breeding-places of, in
Japanese ports, 405, 406;
in Palaearctic Region, 336;
anatomy of, 332.

,, hortensis, in Palaearctic Region,
336 ; anatomy of, 292.

,, (Culiciomyia) impudicus, in
Palaearctic Region, 335.

,, (Lophoceratomyia) jenseni, 336 ;
synonymy of, 78.

5, jugorum (see Aédes pullatus jugo-

yum).

,, laticinctus, in Palaearctic Region,
342.

, laurentit, in Palaearctic Region,
344.

,, lazarensis (see Aédes communis).
,, leucomelas, probably a synonym
of Aédes communis, 314.
,, luteus, perhaps a synonym of C.
pipiens, 345.

,, maculiventris (see Aédes dorsalis).

,, meridionalis, perhaps a synonym
of C. pipiens, 345.

», mimeticus, in Palaearctic Region,
337.

,, mimulus, Anophelines associated
with, 338.

,, (Barraudius) modestus, in Palae-
arctic Region, 332; anatomy
of, 332.

,, molestus, perhaps a synonym of
C. pipiens, 345.

,, musicus, perhaps a synonym of
Aédes rusticus, 308.

», nemorosus (see Aédes communis).

nemorvosus var. dorsovittatus (see

A édes sticticus).

,, nicaensis, perhaps a synonym of
Theobaldia annulata, 288.

,, nigripes var. syluae (see Aédes
sticticus).

,, nigritulus (see C. pipiens).

», niveus, 267.

,, orientalis, sp.n., 342; in Japan,
338; breeding-places of, in
Japanese ports, 407, 408.

494

Culex osakensis (see C. pipiens).

,, pallens (see C. pipiens).
(Culiciomyia) pallidithorax, 335.
, pallipes, 267; perhaps a syno-

nym of C. pipiens, 345.
, perexiguus, in Palaearctic Re-
gion, 342; anatomy of, 343,

344.
,, pipiens, breeding-places of, in
Britain? (205) 72075 °427e-eeam

Japan, 407, 408; in Palae-
arctic Region, 345.
,, (Barrvaudius) pusillus, in Palae-
arctic Region, 333; anatomy
Of Soo Ooo ood
», quasigelidus, in Palaearctic Re-
gion, 337 ; synonymy of, 78.
», quasimodestus (see C. pipiens).
,, vufus (see C. pipiens).
,, salus (see C. sitiens).
,, sergentt, perhaps a synonym of
C. apicalis, 336.
,, simpsont, 78.
sinensis, 78;
gion, 337.
» sttvens, 78, 339 :
water, 29.
,, Spathipalpis (see
longiareolata).
,, taeniorhynchus, breeding in salt
water, 31.
,, taylort, n. n., proposed for Leu-
comyta annulivostris, 78.
,, thalassius, breeding in brackish
water, 29.
,, thovacicus, perhaps a synonym of
C. pipiens, 345.
,, tipuliformis, in Palaearctic Re-
gion, 339, 340, 341; venation
of, 285.
,, trvifilatus, 341.
,, tvitaeniorhynchus, breeding-places
of, in Far Eastern ports, 402—
404, 406, 407; in Palaearctic
Region, 339; in Palestine, 116.
,, Univittatus, 342.
,, varioannulatus (see C. pipiens).
,, ventrilloni, synonymy of, 78.
virgatipes, breeding-places of, in
Hong-Kong, 404; in Palae-
arctic Region, 341.
,, vtrvidis, believed to be a Chirono-
mid, 267.
» vutshnu, 77 ;
gion, 339.
,, vulgaris, doubt as to identity of,
267.
Culicada nemorosa
sticticus).
nigvina (see Aédes sticticus).
Culicella, subgenus of Theobaldia, 286,
289.

in Palaearctic Re-
breeding in sea-

Theobaldia

in Palaearctic Re-

salina (see Aédes

GENERAL INDEX.

culicifactes, Anopheles.

Culicini, key to Palaearctic genera of,
281,

Culiciomyia, a subgenus of Culex, 328,

334.
,, « mnebulosa, perhaps a syno-
nym of Culeximpudicus,
Soon
Culicoides, key to Palestine species of,
108.
iy albicans, 110.
ne civcumscriptus, in Palestine,
108, 115.
* (Johannseniella) fulvithorax,
in Africa, 112.
cy guttularis, sp. n., in Pales-
tine, 108, 114.
* newsteadi, sp. n., in Pales-

tine, 108, 113.

+ odiatus, sp. n., in Palestine,
LOS e012:

i odibilis, sp. n., in Palestine,
108, 114.

of pulicaris, 113, 114.

46 pumilus, 109.

- puripennis, sp. n., in Pales-

tine, 108-109.
a susae, in Italy, 112.
tentorius, sp. n., in Pales-
tine, 108, 110-112.

= vitveipennis, sp. n., in Pales-
tine, 108.
Culiseta siberiensis (see Theobaldia
alaskaensis).
cuniculi, Cuterebra.

curculionis, Thaumasura.
Currant, new Coccid on, in Japan, 216.
curriet, Culex (see Aédes dorsalis).
curtipalpis, Lophoceratomyia (see Culex
Jensent).
Cuterebra_ spp.,
America, 249.
Cyathomyia brevipalpis, referred to
subgenus Neoculex, 79;
(see Culex).
jensen (see Culex).
Cycas vevoluta, Coccid on, in Formosa,
2A:
Cydia pomonella, on peaches in Mauri-
tius, 190.
Cylas formicarius, on sweet potato in
Mauritius, 184.
Cynomyia, characters of larvae of, 258.
Mf cadaverina, Causing Myiasis,
250 ; larva of, 255.
cyprius, Aédes (see A. lutescens).

infesting rodents in

Dacus sygmoides, on cucurbitaceous
plants in Mauritius, 186.

daliensis, Drepanotermes.

danica, Locusta.

GENERAL INDEX,

darwinensis, Mastotermes.
dasypoda, Rogenhofera.
decorella, Tachardia.
Deer, Oestrids infesting, 248, 250.
Deilephila livornica, on grape vines in
Mesopotamia, 479.
Deinocerites, 328.
demodocus, Papilio.
dendrophila, Aédes (Stegomyia).
Dermatobia hominis, causing myiasis,
241, 249.
desertorum, Hypoderma.
designatus, Tabanus (see T. rufino-
tatus).
determinata, Musca (see M. nebulo).
detritus, Aédes (Ochlevotatus).
diana, Hypoderma.
diantaeus, Aédes (Ochlerotatus).
Diaspis, 218.
x boisduvalt, on
Mauritius, 187.
flacourtiae, on Flacourtia in
Seychelles, 128.
7 (Aulacaspis) pentagona, 128 ;
food-plant of, in Formosa,
213; on peaches in Mauri-
tius, 190.
Diatraea sacchaviphaga (see Proceras).
dilatata, Chionaspis.
Dilophus febrilis, 232 ; Gregarine para-
site of larva of, 231.
Dinoderus minutus, in stored maize in
Mauritius, 184.
Dinomimetes, 328.
dipterocarpi, Alcides.
Dipterocarpus tuberculatus, new weevil
bred from seeds of, in India, 167.
disjunctus, Oestrus.
dispar, Heterogamus.
divergens, Simulium.
Diversinervus stlvestrvii, parasite of
Coccus viridis in Mauritius, 188.
diversipes, Rhadinomerus.
Doctostaurvus maroccanus, 159, note.
domestica, Musca.
domesticus, Culex.
Donkeys, trypanosomiasis in, in Kenya
Colony, 234.
dorsalis, Aédes (Ochlevotatus).
dorsomaculata, Musca (see M. albo-
maculata).
dorsovittatus, Culex
Aédes sticticus).
Drepanotermes daliensis, sp. n., in Aus-
tralia, 373-375.

coconut in

nemorosus (see

i pernigey, in Australia,
374-375.

x rubriceps, 364, 367, 373,
376-378.

septentrionalis, sp. n.,
bionomics of, in Aus-
tralia, 368, 372.

495

Drepanotermes silvestrii, sp. n., biono-
mics of, in Australia, 364-371.

Drosophila, larva of, accidentally pro-
ducing myiasis, 252.

dubium, Simulium hirsutum.

duplex, Pseudaonidia.

Dutch East Indies, mosquitos in, 70.

dux, Chrysomyia (see C. megacephala).

dyart, Theobaldia.

Dysdercus, on cotton in
132.

Dysis, 474.

Mauritius,

Earias insulana, on cotton in Mauritius,
182.

eatoni, Aédes (Finlaya).

echinus, Aédes (Finlaya).

Ecculex, subgenus of Aédes, 293, 322.

egregia, Osphilia.

Egypt, Hippobosca camelina on camels
in, 122 ; mosquitos in, 264.

Elephant, Oestrids infesting, 247, 248,
249.

elephantis, Cobboldia.

elestéem, Tabanus (see T. rufinotatus).

Elk, Oestrid infesting, 250.

elongatus, Coccus.

elutus, Anopheles.

emasculator, Bogeria.

Ephestia cautella, on ground-nuts in
Mauritius, 186.

ephippium, Cuterebra.

Epipyrops fuliginosa, sp. n., parasi-
tising Idiocerus in India, 466, 468.
equi, Gastrophilus (see G. intestinalis).

equina, Hippobosca.
equinum, Simulium.
Eycta ornatalis, on sweet potato in
Mauritius, 184.
Eriococcus graminis, Antonina crawtt
recorded as, 211.
Eristalis, characters of larva of, 258.
is tenax, accidentally producing
myiasis 252.
erythrocephala, Calliphora.

Ethiopian Region, mosquitos from,
found in Palaearctic Region, 264.
Eucalymnatus tessellatus, food-plants of,

in Mauritius, 185, 189.
eucalypti, Hamitermes.
Eucalyptus, termites

Australia, 394.
Eugenia, new weevils bred from, in

India, 175, 177.
PF caryophyliata, new scale on,
in Seychelles, 125.
eugeniae, Chionaspis ; Phenacaspis.
euonymi, Aphis (see A. rumicis).
Euonymus, food-plant of Aphis rumicis,
81, 86, 87.

infesting, in

496

Eupristina verticillata, sp. n., from figs
in Borneo, 38-40.

Europe, representative mosquitos of,
265.

eusivus, Microplitis.

eutaeniata, Musca (see M. humilis).

Eutermes, not attacked by Ividomyr-

mex, 371.
. tyviei, in Australia, 386.
euthyrrhini, Chalcis.

Euthyrrhinus meditabundus, bionomics
of, on mangos in Australia, 63-66 ;
new parasite of, 67.

excrucians, Aédes (Ochlerotatus).

fabae, Aphis (see A. rumicis).
falciparum, Plasmodium.
Fannia, larva of, accidentally pro-
ducing myiasis, 252.
canicularis, causing myiasis,
241; found in bladder of
man 250: characters of
larva of, 258, 260:
a scalaris, characters of larva of,
258, 260.
Far East, mosquitos found in ports of,
401-409.
fasciata, Bogeria ; Stegomyia (see A édes
argenteus). (
fasciatus, Culex.
fatigans, Culex.
febrilis, Dilophus.
Federated Malay States, new mosquitos
in, 72, 79 ; mosquito larvae in, 91-97.
fergusoni, Musca (see M. convexifrons).
fevox, Leucotermes.
ferruginea, A phiochaeta
xanthina).
ficalbi, Culex (see Theobaldia fumi-
pennis).
ficoides, Cerococcus.
Ficus spp., fig insects from, in Borneo,

(see 4.

Ficus retusa, Coccid on, in Japan, 212.

ficus, Chrysomphalus.

Fig Insects, new, from Borneo, 35-40.

Figs, bionomics of Ocnerogyia amanda
on, in Mesopotamia, 477.

Fiji, new mosquito in, 79.

filamentosus, Pseudococcus.

filipes, Rachisoura (Culex).

finalis, Tabanus.

Finland, new mosquitos in, 306, 314.
Finlaya, subgenus of Aédes, 293, 317 ;
key to Palaearctic species of, 317.
Fiorinia chinensis, sp. n., in China,

215, 216.

a froriniae on palm in U.S.A.,,
Zo:

Re japonica, food-plants of, in
Formosa, 215.

GENERAL

INDEX.

Fiorinia juniperi, food-plants of, in
Formosa, 215.
* yvubrolineata, 216.
froriniae, Fiorinia.
Flacourtia, Diaspis flacourtiae on, in
Seychelles, 128.
flacourtiae, Diaspis (Aulacaspis).
flava, Cosmophila. °
flaviceps, Anopheles ; Chrysomyia.
flavipes, Gastrophilus ; Simulium.
flavivostris, Culex.
flaviveniris, Leptoconops.
flavocincta, Chrysops.
flavovirens, Culex.
Flax, Lepidopterous pests of, in Kenya
Colony, 99-102.
flerowi, Anopheles pseudopictus.
fontenella, Bogeria.
Forcipomyia bipunctata, in Palestine,
115.
fovestan, Rhopalocampta.
formicarius, Cylas.
Formosa, Coccids in, 211-220.
foveicollis, Agrilus.
fraseri, Aédes (Stegomyia) ; Leptosoma-
tomyia.
frater, Sympiezomias.
freyi, Aédes (Ochlerotatus).
Fuchow, breeding-places of mosquitos ~
at, 40).
fuliginosa, Epipyrops.
fuliginosus, Anopheles ; Silvius (see S.
notatus).
fulvicollis, Plecia.
fulvimedioides, Tabanus.
fulvimedius, Tabanus.
fulvithovax, Culicoides (Johannseniella).
fumipennis, Theobaldia (Culicella).
fumosus, Mecistocerus.
funebris, Cuterebra.
funerea, Aédes (Skusea).
fusculus, Culex (see Aédes geniculatus).
fuscus, Culex (Neoculex, Protomelano-
conion) ; Optfex.

Galinosoga parviflora, food-plant of
Phytometra orichalcea, 99.
gallit, Aédes (see A. pullatus jugorum).
Gardenia florida,Coccid on, in Japan, 212.
Gasterophilus (see Gastrophilus).
Gastrimargus musicus, 162.
Gastrophilus equi (see G. intestinalis).
ms flavipes, distribution of,
245.
- gedoelstt, infesting zebra,
245.
a4 haemorrhoidalis, in horses,
distribution of. 244.
= intestinalis, in horses, dis-
tribution of, 244; larva
of, 261.

GENERAL INDEX.

Gastrophilus intestinalis var. asininus,
244.

intestinalis var. bengalensis,
244; egg of, 261.

lativentris, in Courland,
245.

magnicornis, possibly iden-
tical with G. intestinalis
var. bengalensts, 245.

nasalis (see G. veterinus).

nigrvicollis, in Bessarabia,
245.

af pecorum, in horses,
tribution of, 245.

pecovrum zebrae, infesting

dis-

zebra, 245.
~ ternicinctus, infesting zeb-
Tas 240%
veteyinus, in horses, dis-

tribution of, 245; larva
of, 261.

gazellae, Hypoderma.

gedoelsti, Gastrophilus.

Gedoelstia cristata, infesting antelopes
in Africa, 247.

hassleri, infesting antelopes
in Africa, 247.

Gelastorrhinus sagitta, breeding-places
of, in Transcaspia, 154.

geniculatus, Aédes (Finlaya).

gerymanus, Hamitermes.

Germany, Bibionid larvae attacking
potatoes in, 232.

germinatus, Mandalotus.

gibsomt, Musca.

glaphyroptera, Theobaldia.

globulus, Tvigonogenius.

glovert, Lepidosaphes.

Glugea, infesting Bibionid larvae, 231.

Glyphodes indica, on cucurbitaceous
plants in Mauritius, 186.

Goats, Lipoptena caprina on, in Pales-
tine, 124.

Gold Coast, new mosquito in, 74.

gossypiella, Platyedra.

gossypit, Aphis.

Gossypium herbaceum, Coccid on, in
Japan, 212.

Grabhamia broquettii(see A édes dorsalis).

grahami, Aédes (see A. dorsalis).

graminis, Eviococcus.

grandis, Leptoconops ; Rogenhofera.

gvavelyi, Megarhinus (Toxorhynchites).

grisea, Bogeria.

grossipes, Cryptodus.

Ground-Nut (Avachis hypogaea), pests
of, in Mauritius, 186.

Guava (Psidium), scale-insects on, in
Mauritius, 182.

gurneyi, Philaematomyia (see Musca
(Ptilolepis) inferior).

guttularis, Culicoides.

”

497

Gyrostigma, larva of, 254.

xs conjugens, infesting rhino-
ceros in Africa, 245.

Pe meruensis, infesting rhino-
ceros in Africa, 245.

* paves, infesting rhino-
ceros in Africa, 245.

Bs sumatrensis, infesting rhi-
noceros in East Indies,
245.

Haartebeeste, Oecestrids
Africa, 246, 247.

hackert, Anopheles leucosphyrus.

Hadronotus aniestiae, parasite of An-
testia lineaticollis in Kenya Colony,
191—201.

Haematopota alluaudi, attacking cattle

infesting, in

in Kenya Colony,
xe Toys:

= brunnescens, attacking
cattle in Kenya

Colony, 235.
* cilipes, in Siam, 432.
hirta, attacking cattle in
Kenya Colony, 235.
7 pachycera, in Siam, 432.
: similis, attacking cattle
in Kenya Colony, 235.
- ugandae, attacking cattle
in Kenya Colony, 235.

a validicornts (see H. pachy-
ceva).
Haemoproteus columbae, carried by

Lynchia maura, 122.
haemorrvhoidalis, Gastrophilus.

halifaxi, Lutzia.

Hamitermes eucaly pti, sp.n., bionomics

of, in) Australias 385,
391-395.
Re germanus, 383.

herbertensis, 393.

latidens, 381.

laurensis, 384, 386.

mervidionalis, 384, 386.

¥ neogeymanus, sp. n., in
Australia, 390.

obtusidens, 382.

parvus, sp. n., in Australia,
378-381.

_ per plexus, sp.n., bionomics
of, my, Australia, 371,
381-388, 390, 395.

perplexus var. victoriensis,
n., in Australia, 388.

Harpagomyia, 282.

hasslevi, Gedoelstia.

hayashi, Culex.

Heliothis (Chloridea) obsoleta, on flax
in Kenya Colony, 99; food-plants
of, in Mauritius, 182-186.

”?

498

Helophilus, characters of larva of, 258.

pendulus, accidentally pro-
ducing myiasis, 252.

Hemichionaspis, considered a synonym
of Pinnaspis, 214.

Hemileia vastatrix, on coffee in Mauri-
tius, 187.

hemisphaerica, Saissetia.

Henicospilus antancarus, parasite of
Sesamia vuteria in Mauritius, 184.

herbertensis, Hamitermes.

hermanii, Pseudogametes.

Herse convolvuli, on sweet potato in
Mauritius, 184.

hesperidum, Coccus.

Heterodeva vadicicola, food-plants of, in
Mauritius, 183, 186.

Heterogamus, treated as a subgenus of

Rhogas, 130.
- dispar, 130, 132.

hewitti, Cevatosolen.

Hibiscus esculentus, food-plant of
Chloridea obsoleta in Mauri-
tius, 182.

yosa-sinensis, Coccid on, in
Japan, 212.

hilavis, Tabanus (see T. tenens).

hilli, Mimeteomyia (see Fachisoura
filipes) ; Musca (see M. ventrosa).

Hippobosca camelina, in Palestine,
122:

capensis, in Palestine, 122.

a. equina, in Palestine, 121.

hippopotami, Rhinoestrus.

Hippopotamus, Oestrid infesting, in
Africa, 247.

hirsuteron, Aédes.

hirsutum, Simulium.

hirta, Haematopota.

hispaniola, Anopheles (Myzomyia).

histrio, Cuterebra.

Hodgesia, 282.

Hollyhock, food-plant of Asterochiton
vaporariorum in Britain, 359.

Holoconops, a subgenus of Leptoconops,
oF 2 LOd. Wore:

hololeucus, Niptus.

hominis, Dermatobia.

Horses, Oestrids infesting, 244, 245 ;
Simulium attacking, in Palestine,
117.

hortensis, Culex.

hortulanus, Bibio.

humilis, Musca (see M. pumila).

hyalinipennis, Leptoconops (see L.
bezzit).

Hypocera incrassata, parasite of larva
of Bibio marci, 231.

Hypoderma, characters of larva of, 255.

actaeon, infesting red deer,
248.

»”

”?

»”?

GENERAL INDEX.

Hypoderma bovis, distribution of, on

cattle, 248; causing
myiasis, 241, 248; larva
(one, PAS.

corinnae, infesting gazelles
in Africa, 248.

desevtorum, in Egypt, 248.

diana, infesting red deer,
248.

gazellae, infesting gazelles
in Africa, 248.

lineata, distribution of, on
cattle, 248; larva of,
261.

silenus, infesting donkeys
in Egypt, 248.

hyvcanus, Anopheles.

”

ibicis, Lipoptena.

Icervya purchasi, food-plant of, in For-
mosa, 211) on cltrs sand
coffee in Kenya Colony, 103.

seychellarum, food-plants of, in
Formosa, 211; food-plants
of,in Mauritius, 185, 188,189.

Idiocerus spp., natural enemies of, in
India, 465-469.

Ilex crenata, new Coccid on, in Japan,
216.

immaculatus, Anopheles.

impudicus, Culex (Culiciomyia).

incrassata, Hypocera.

indefinitus, Anopheles rossi
subpictus var. vagus).
indemnis, Bubaris.

India, new Braconidae in, 129-132 ;
flies causing myiasis in, 239 ; house-
flies in, 415-423; natural enemies
of mango leaf-hoppers in, 465—469 ;
new mosquitos in, 72, 73, 77; new
weevils in, 165-180.

indianus, Tabanus.

indica, Glyphodes; Musca (Philaema-
tomyia) (see M. cingalaisina).

indicus, Leptoconops.

inferior, Musca.

infulata, Cuterebra.

innuitus, Aédes (see A. alpinus).

insidiator, Tabanus.

insignis, Musca (Philaematomyia) (see
M. crassirostris).

insulana, Eartas.

interruptus, Leptoconops ; Oestrus.

intestinalis, Gastrophilus.

intrudens, Aédes (Ochlerotatus).

inusitatus, Aspidiotus.

tota, Pseudaonidia.

Iridomyrmex sanguineus, relation of, to
termites in Australia, 371, 388.

irvitans, Leptoconops ; Lyperosia.

Italy, Culicoides susae in, 112.

ce)

(see A.

GENERAL INDEX.

jacobsont, Blastophaga.
janata, Achaea.
Janthinosoma lutzi, carrying eggs of

Dermatobia hominis,
249,
ms musica (see Psoro-

phora posticata).
Japan, Coccids in, 211, 215-217; new
mosquitos in, 274, 327, 338; mos-
quitgs found in ports of, 401-409.

japonica, Ftorinia; Lepidosaphes ;
Mytilaspis pomorum (see Lepido-
saphes).

japonicus, Aédes (Finlaya).

Java, Coccids in, 212, 213.

gensent, Culex (Cyathomyia, Lophocera-
tomyia).

johannis, Bibio.

Johannseniella fulvithovax
coides).

jugorum, Aédes pullatus (Culex).

junipert, Fiorinia.

(see Culi-

harwart, Anopheles.

kasauliensis, Musca (see M. ventrosa).

Kashmir, house-flies in, 423.

kempi, Megarhinus (Toxorhynchites).

Kenya Colony, bionomics of Chry-
somphalus aurantit in, 103-104 ;
bionomics of parasites of the coffee
bug in, 191-201 ; trypanosomiasis in
the absence of Glossina in, 233-236 ;
Lepidopterous pests of flax in, 99—
102 ; human disease possibly carried
by Simulium in, 236-238.

kertész1, Leptoconops.

Kirkioestrus spp., infesting antelopes
in Africa, 247.

klossi, Megarhinus (Toxorhynchites).

Kobe, breeding-places of mosquitos
at, 406.

kochi, Anopheles.

koreicus, Aédes (Finlaya).

Lachnosterna smithi, food-plants of, in
Mauritius, 184, 185.

lacinia, Pseudaonidia.

lacteipennis, Bibio; Leptoconops.

lacteus, Coptotermes.

Laingia psammae, gen. et sp. n., on
grasses in Britain, 429.

lambens, Sarcophaga.

Lam pides boetica, on Cajanus indicus in
Mauritius, 185.

lapatht, Cryptorrhynchus.

larvipara, Musca.

lataniae, Aspidiotus.

laticinctus, Culex.

latidens, Hamitermes.

latiparafrons, Musca (see M. gibsont).

499

lativentris, Gastrophilus.
laurae, Leptoconops (see L. kertész1).
lauvensis, Hamitermes.
laurventi, Culex.
lazavensis, Aédes.
leicester1, Megarhinus.
Leicesteria annulipalpis, in Sumatra,
74.
lepidonotus, Aédes (Ochlerotatus).
Lepidosaphes beckit, food-plant of, in
Formosa, 216.
a gloveri, on citrus in Mau-
ritius, 188.
Ff japonica, food-plant of, in
Japan, 217.
= tubulorum, sp. n., food-
plant of, in Formosa
and Japan, 216.
aA ulmi, L. tubulorum re-
corded as, 217.
Lepidotomyia lineata (see Aédes funerea
var. ornata).
Leptoconops, revision of genus, 1-28 ;
key to females of, 27.
me bezzii, 2; anatomy of, 19;
description of, 17.
a braziliensis, anatomy of,
8, 19; description of, 13.
is flaviventris, description of,

18.

5 grandis, sp. n., anatomy
of, 9; 195. description
of, 12.

hyalinipennis (see L. bez-
Z11).

7 indicus, description of, 19.

- interruptus, 23.

ie irvitans, bionomics of, 3;
description of, 14 ; not
experimentally infected
with malaria, 4.

mys hkertészi, bionomics of, 4 ;
anatomy of, 19; des-

cription / ‘of, 21; 4m
Palestine, 107.
“A kervlészi var. americanus,

n., anatomy of, 5-9, 19 ;
description of, 22.
‘5 kertészi var. peneti, 6;
description of, 22.
es lacteipennis, anatomy of,
9; description of, 23.
laurae (see L. kertészt).
longicornis, sp. n., anat-
omy of, 9, 19; descrip-

tion of, 11.

a vhodesiensis, sp. n., anat-
omy of, 6, 9; descrip-
tion of, 14.

re siamensis, Sp. n., anatomy
of, 6, 8, 9, 19; descrip-
tion of, 20.

500

Leptoconops spinosipes (see Acantho-
conops albiventris).

stygius, 1; description of,
10; anatomy of, 6, 8,

”

O19:
as torvens, 1, 15-17; anatomy
of, 6, 8, 9, 19.

Leptosomatomyta fraseri, 286.

lesnet, Ochlerotatus.

leucocnematus, Tabanus (Atylotus).

leucomelas, Culex.

Leucomyia annulata, new name sug-
gested for, 78, note.

annulirostris (see Culex tay-
lort).

leucosphyrus, Anopheles.

Leucotermes clarki, sp. n., in Australia,
395-398.

ferox, in Australia, 398.

pavadoxus, in Australia,
363.

Ss validus, in Australia, 398.
lewisi, Anopheles (see A. maculipennis).
Lime (Citrus medica var. acida), pests

of, in Mauritius, 187.
lindesayi, Anopheles.
lineata, Hypoderma ; Lepidotomyia (see
Aédes funerea var. ornata) ; Musca
(Pristivhynchomyia).
lineaticollis, Antestia.
lineatum, Simulium (see S. maculatum) .
lineatus, Tabanus (see T. rufinotatus).
Lipoptena caprina, sp. n., on goats in
Palestine, 122, 124.
cervt, 124.
chalcomelaena, on goats in
Palestine, 124.
ibicis, possibly a synonym of
L. chalcomelaena, 124.
Litchi (Nephelium), scale-insects on, in
Mauritius, 182.
litura, Prodenia.
livornica, Deilephila.
Locusta, revision of genus, 135-163.

a (Pachytylus) brasiliensis (see
Locusta migratoria ph. da-
nica).

capensis (see Locustana par-
dalina).
danica, considered to be a
phase of L. migratoria, 137.
migratoria, bionomics and mi-
grations of forms of, 135-163;
dimensions of phases of,
139 ; key to phases of, 161.
migratorioides, considered to be
a phase of L. migratoria, 150.
,, pardalina (see Locustana).
,, sulcicollis (see Locustana par-
dalina).
Locustana, gen. n., erected for Locusta
pardalina, 137, 162.

”

GENERAL INDEX.

Locustana pardalina, bionomics of, in

South Africa, 155-159 ;
key to phases of, 163.

Pe pardalina solitaria, ph. n.,
163.

Locusts, biology and migrations of,
143-148.
longiarveolata, Theobaldia (Allotheobal-
dia).
longicornis, Leptoconops.
Lophoceratomyia, a subgenus of Culex,
328 ; Cyathomyiaa
synonym of, 79.
ae curtipalpis (see Culex
jensen).
loxodontis, Cobboldia.
Lucilia, characters of larvae of, 258,
260.
sf argyvricephala, causing myiasis,
289Fe 250 larva Ole o 4.
2595.
,, eaesay, Causing myiasis, 250.

» sevenissima (see L. argyrice-
phala).

a sevicata, causing myiasis, 240,
25) latvias Ot e204 OOF
Jove

By tasmaniensis, Causing myiasis,
250.

ludlow1, Anopheles.
lukist, Anopheles (see A. algeriensis).
lunata, Chilomenes.
lusoria, Musca.
lutescens, Aédes (Ochlerotatus).
luteus, Culex.
lutzi, Janthinosoma.
Lutzia concolor, 327.
,, halifax, 327.
,, tigripes, 337. ;
|, vovax, Sp. T., in” Japan, 3270:
breeding-places of, in Japan-
ese ports, 405-406 ; anatomy
of, 285, 292.
Lynchia maura, on pigeons in Palestine,
122.
Lyperosia irritans, in Palestine, 121.
* minuta, in Palestine, 121.
lysimon, Zizera.

macdonaldi, Oestrus.

macellavia, Cochliomyia (Chrysomyia).

Macrosiphum picridis, on artichoke in
Mauritius, 186.

macrotis, Cephenomyia.

maculata, Cephalomyia (see Cephalopsis
titillator) .

maculatum, Simulium.

maculatus, Aédes (Ochlerotatus) ;
pheles.

maculipalpis, Anopheles.

maculipennis, Anopheles ; Plutella.

Ano-

GENERAL INDEX.

maculiventris, Culex Aédes
dorsalis).
maculosa, Cuterebra.
magnicornis, Gastrophilus.
magnifica, Wohlfahrtia.
matdis, Aphis.
Maize, pests of, in Mauritius, 183.
Malaria, Leptoconops irritans not ex-
perimentally infected with, 4; in-
digenous cases of, in England, 209.
malefactor, Anopheles.
Mallococcus sinensis, in China, 212.
malloti, Rhadinomerus.
Mallotus japonica, Coccid on, in Japan,
D2:
e philippinensis, new weevil
bred from, in India, 176.
Man, Diptera producing myiasis in,
239-261; disease of, probably con-
veyed by Simulium, 236-238.
Mandalotus geyminatus, in termite
nests, 371.
mandersi, Nacaduba.
mangiferae, Coccus; Cryptorrhynchus.
Mango, Euthyrrhinus meditabundus and
termites on, in Australia,
63-66, 371; pests of, in
Mauritius, 182, 189.
Mango Leaf-hopper (see Idiocerus).
Manioc (Manihot utilissima), pests of,
in Mauritius, 184.
Mansonioides, subgenus of Taeniorhyn-
chus, 291.
marci, Bibio.
marginale, Chrysomyia.
mariae, Aédes (Ochlerotatus).
marocanus, Culex annulatus (see Theo-
baldia longiareolata).
maroccanus, Dociostaurus.
mashonaensis, Uvanotaenia.
Mastotermes darwinensis, attacking
mango, etc., in Australia, 63, 371.
matteiana, Procontarinia.
maura, Lynchia.
mauritia, Spodoptera.
mauritianus, Anopheles.
mauritit, Stauropodoctonus.
Mauritius, Coccids in, 213; pests of
minor crops in, 181-190.
Mecistocerus, Rhadinomerus compared
with, 170.
f fumosus, sp. n., on Pinus
longifolia in India, 168-
0:
meditabundus, Euthyrrhinus.
megacephala, Chrysomyia; Pheidole.
megalops, Tabanus (see T. tenens).
Megarhinus brevipalpis, 285.
Ae christophi, in Palaearctic
Region, 285.
ri (Toxorhynchites) gravelyi,
Spee in India. "73:

(see

S01

Megarhinus (Toxorhynchites) hemi, sp.
mine India “72:
A (Toxorhynchites) klossi, sp.
n., in Federated Malay
States, 72.

y leicesteri, 72.

* vegius, 285.

We vutilus, 285.

Be towadensis, in Palaearctic

Region, 285.
megastoma, Cuterebra.

meigenanus, Aédes (Ochlerotatus) punc-

tor.

mella, Tachina.

Melons, pests of, in Mauritius, 186.

meridionalis, Culex ; Hamitermes.

meruensis, Gyvostigma.

Mesopotamia, Lepidopterous pests in,

477-479 ; mosquitos in, 264.
mesopotamiae, Anopheles hyrcanus.
metalepticus, Aédes (see A. pullatus

jugorum).

Michelia fuscata, Coccids on, in For-

IMOSaw 2S -220).

Micraédes, a subgenus of

328.

Microplitis eusivus, sp. n., parasite of

Achaea janata in India,
129:

ee similis, sp. n., parasite of
Agrotis ypsilon in India,
129.

-f spectabilis, 129.
microptera, Stegomyia.
migratoria, Locusta.
migratorioides, Locusta.
Mimeteomyia hilli (see

filipes).

mimeticus, Culex.

mimulus, Culex.

minimus, Anopheles.

minor, Musca (see M. pumila); Pin-

naspis ; Pternoscirta (Pachytylus).
minuta, Lyperosia ; Musca.

minutus, Dinoderus ; Kirkioestrus.

miscanthi, Pygalataspis.

Miscanthus sinensis, new Coccid on, in

Formosa, 218.

modestus, Culex (Barvaudius).

molestus, Culex.

moluccensis, Anopheles punctulatus.

montanus, Anopheles albotaeniatus.

morio, Alcides.

morsitans, Theobaldia (Culicella).

Morus alba, Coccids on, in Japan, 211,

213,219;

Mosquito Larvae, function of caudal
tufts of, 91-97; in saline
waters, 29-34; destroyed
by Tabanid larvae, 22.

= Larvicides, 32.

Culex,

Rachisoura

502

Mosquitos, classification and new
species of, 69; breeding-places of, in
Britain, 205-209, 427; of Far East-
ern ports, 401-409 ; from the Palae-
arctic Region, 263-351; keys to,
269, 276, 281, 286, 293-295, 296-299,
317, 324, 329-331; bibliography of
Palaearctic, 348; index of specific
and varietal names of, 350.

mucronata, Schneideria.

multicolor, Anopheles (Myzomyia).

Murraya exotica, Coccids on, in Japan,
QAZ 216:

Musca, notes on genus, 411—426.

,, albina, 423.
,, albomaculata,
420.

,, angustifrons (see M. humilis).
,, australis (see M. convexifrons).
,, autumnalis, 121; bionomics of,

in England, 421.
, vbezzit, in India, 421.
,, bivittata (see M. humilis).

distribution of,

,, eingalaisina, distribution of,
422.

,, econducens (see M. humilis).

,, convexifrons, in Australasia,
425.

,, corvina (see M. pumila).

,, (Philaematomyia) crassivostrts,

in Palestine, 120 ; characters
of larva of, 254 ; distribution
of, 423.

,, deteryminata (see M. nebulo).

., domestica, 121; atypical form
of, in Australasia, 424;
atypical form of, in India,
416; list of synonyms of,
418; discussion as to world-
wide distribution of, 411—
414; characters of larva of,
254,257,298:

,, dorsomaculata (see M.
maculata).

,, eutaeniata (see M. humilis).

, fergusoni (see M. convextfrons).

,, gibsont, in India, 420.

» Arllt (see M. ventrosa).

,, humilis, bionomics and distri-
bution of, 417-419; syno-
nymy of, 417; characters of
larva of, 254.

», (Philaematomyia) indica (see M.
cingalaisina).

,, tmferior, in India, 422.

,, (Philaematomyvia) imsignis (see
M. crassivostris).

,, Rasauliensis (see M. ventrosa).

», larvipara, 421.

,, latiparafrons (see M. gibson).

,, lineata, in India, 422.

» lusoria, 425.

albo-

GENERAL INDEX.

Musca minor (see M. pumila).
3 6 6©minuta, 423.
,, nebulo, an Oriental species, 414,
415; characters of larva of,
254, 257.
» negriabdomina, 423.
», nigrithorax (see M. ventrosa).
», niveisquama (see M. humilis).
,, pattoni, in India, 420.
, ptlosa (see M. bezzit).
,, pollinosa (see M. cingalatsina).
», praecox (see M. humilis).
» primitiva (see M. humilis).
» promisca (see M. humilis).
,, pumila, in Australasia, 424 ; in
India, 419.
,, pungoana (see M. ventrosa).
, setigera (see M. albomaculata).
,, Sspeculifera, 423.
,, Spinohumera, bionomics of, in
India, 421.
,, Spinosa (see M. pattonz).
,, . striatecta, 423.
,, ltempestiva, in Kashmir, 423.
,, terrvae-veginae, in Australasia,
425.
,, ventyosa, in Australasia, 425;
in India, 419.
,, vetustissima (see M. pumila).
,, vicaria (see M. domestica).
,, vttyipennis, in Kashmir, 423.
,, «xanthomela (see M. ventrosa).
musica, Janthimosoma (see Psorophora
posticata).
musicus, Culex ; Gastrimargus.
Mycterotypus, 1; synonym of Lepto-
conops, 3.
Myiasis, Diptera producing, in man
and animals, 239-261.
Myrica rubra, Coccid on, in Japan, 212.
Mytilaspis pomorum (see Lepidosaphes
ult).
-s pomorum var. japonica (see
Lepidosaphes japonica).
Myzomyia, subgenus of Anopheles, 275.

Nacaduba mandersi, on Cajanus indicus
in Mauritius, 185.

Nagasaki, breeding-places of mosquitos
at, 404.

nasalis, Gastrophilus (see G. veterinus).

nearcticus, Aédes (Ochlerotatus) (see A.
alpinus).

neavet, Simulium.

nebulo, Musca.

nebulosa, Culiciomyia.

negriabdomina, Musca.

nemorosus, A édes (Ochlerotatus).

Neoculex (see Culex).

neogermanus, Hamitermes.

nephodes, Tabanus (Atylotus).

GENERAL

New Zealand, Opifex fuscus in, 73, 74.

newsteadi, Culicoides.

nicaensis, Culex.

Nigeria, new Simuliids in, 459-463, .

nigra, Saissetia ; Stomoxvs.

nigricans, Cuterebra.

nigricollis, Cercyonia ; Gastrophilus.

nigrina, Culicada (see Aédes sticticus).

nigritarsis, Tabanus.

migrithorax, Musca (see M. ventrosa).

nigritulus, Culex (see C. pipiens).

nigvocincta, Cuterebra.

nigropunctata, Crioceris.

nigvotectus, Tabanus (Bellardia).

nipponit, Aédes vexans.

Niptus hololeucus, 133.

nivarleti, Rhinoestrus.

niveisquama, Musca (see M. humilis).

nivettaeniata, Theobaldia (Pseudotheo-
baldia).

niveus, Aédes (Finlaya) ;

niveosparsus, Idiocerus.

notata, Scatopse.

notatus, Silvius.

Culex.

obovata, Aspidomorpha.
obscurus, Aédes (seer 241. communis) ;
- Calotermes.
obsoleta, Heliothis (Chloridea).
obturbans, Armigeres.
obtusa, Aonidia.
obtusidens, Hamitermes.
occidentalis, Anopheles (see A. maculi-
pennis).
Ochlerotatus, key to Palaearctic species
of, 296-299 : subgenus
of Aédes, 293, 295 ; g.v.
it annulifemur (see Stegomyia
periskeleta).
Ocnerogyia amanda, bionomics of, on
figs in Mesopotamia, 477.
octoguttalis, Thliptoceras.
octoguttatus, Botys.
odiatus, Culicoides.
odibilis, Culicoides.
Odina wodier, new weevil bred from, in
India, 180.
odinae, Osphilia.
Odonaspis, Pygalataspis resembling,218.
penicillata, food-plant of,
in Japan, 219.
Oedemagena tavandi, infesting reindeer
in Scandinavia, 248.
Oestrus aureo-argentatus, infesting ante-
lopes in Africa, 247.
», compositus, infesting antelopes
IMpAiTICa 247
,, aisjunctus, infesting antelopes
in Africa, 247.
», interruptus, infesting antelopes
in Attica 247).

a»

(6127)

INDEX. 503

Oestrus macdonaldi, infesting antelopes
in Africa, 247.
», ovis, infesting Sheep, 244, 246 ;
characters of larva of, 255.
», vartolus, infesting haartebeeste,
246.
oleae, Argopistes ; Saissetia.
Olives, new Coleopterous pests of, in
South Africa, 474, 475.
Opifex fuscus, considered to be a
Culicine, 73 ; description of, 73, 74.
Opuntia, extract of, as an adhesive in
insecticides, 185.
orichalcea, Phytometra.
Oriental Region, mosquitos from, found
in Palaearctic Region, 264.
orientalis, Culex.
ornata, Aédes (Skusea) funerea.
ornatalis, Eycta.
Orthopodomyia albionensis (see O. pul-
chripalpis).
i pulchripalpis, in Palae-
arctic Region, 290.
Oryctes tavandus, bionomics of, on
coconut in Mauritius, 187.
oryzae, Calandra.
osakensis, Culex (see C. pipiens).
Osphilia brevirostris, in Ceylon, 180.
Bi egvegia, in Burma, 180.
nh odinae, sp.n., bred from forest
trees in India, 179.
ovis, Oestrus.
Oxya tuvanica, breeding-places of, in
Transcaspia, 154.

pachyceva, Haematopota.
Pachytylus (see Locusta).
i austvalis, a Synonym of
Locusta migratoria ph.
danica, 162.
ae capito, a synonym of Lo-
custa migratoria ph. mi-
gvatorioides, 162.
er minor (see Pternoscirta).
Palaearctic Region, mosquitos of, 263-
351.
Palaeopsyche, 468.
Palestine, blood-sucking Diptera of,
107-124 ; mosquitos in, 264.
palestinensis, Anopheles (Pyretophorus)
(see A. superpictus).
pallens, Culex (see C. pipiens).
pallidothorax, Culiciomyia.
pallipes, Culex.
palment, Aédes (Ochlerotatus) (seeme
communis).
palmert, Simulium.
paludis, Anopheles (see A. mauritianus)
panayensis, Aédes.
Pandanus seychellarum, Pinnaspis buxi
on, in Seychelles, 127.

2102

504

papatasii, Phlebotomus.

papaveris, Aphis (see A. rumicis).

Papilio demodocus, on citrus in Mauri-
tius, 188.

phorbanta, on citrus in Mauri-
tius, 188.

paradoxus, Leucotermes.

parcus, Rhadinopus.

pardalina, Locustana (Locusta).

Parlatoria pergandet, food-plants of,
in Formosa, 214.

zizyphi, food-plant of, in
Formosa, 214.

partitus, Tabanus (see T. striatus).

parumspinosa, Cobboldia.

parvulus, Aédes (Ochlerotatus).

parvus, Hamitermes.

patagona, Cuterebra.

pattont, Musca.

paves, Gyrostigma.

Peaches, pests of, in Mauritius, 190.

Peas (Pisum sativum), pests of, in
Mauritius, 186.

pecorum, Gastrophilus.

Pelamia repanda (see Remigia).

pembaensis, Aédes (Skusea).

pendulus, Helophilus.

peneti, Leptoconops kertészt.

penicillata, Odonaspis.

pentagona, Diaspis (Aulacaspis).

percurrens, Rhogas (Heterogamus) .

peregrina, Schistocerca.

perexiguus, Culex.

pergandet, Parlatoria.

periskeleta, Stegomyia.

perniger, Drepanotermes.

perplexus, Hamitermes.

persicus, Anopheles (Myzomyia) turk-

hudi.

pfeiffert, Xanthogramma.

phacochoert, Rhinoestrus.

pharoensis, Anopheles (Myzomyia).

phaseoli, Agromyza.

Pheidole megacephala, 64.

Phenacaspis eugeniae, food-plant of,
in Formosa, 213.

Philaematomyia, characters of larva of,
258 ; (see Musca).
gurneyi (see Musca

(Ptilolepis) inferior).
Phlebotomus minutus var. africanus, in
Palestine, 119.
papatasit, 115; in Pales-
tine, 118.
phobtifer, Cephenomyia.
phorbanta, Papilio.
Phormia, characters of larva of, 258.
regina, Causing myiasis, 250 ;
larva of, 255:
Phragmites communis, locusts breeding
amongst, in Transcaspia, 143, 154.
Phytalus smithi (see Lachnosterna).

»”

»>

”

GENERAL INDEX.

Phytometra chalcytes, food-plants of,
in Mauritius, 183.

i ovichalcea, bionomics and
control of, on flax in
Kenya Colony, 99-102;
food-plants of, in Mau-
ritius, 183, 186.

Phytophaga, new injurious species of,
from, Atricay 473-475:

picridis, Macrosiphum.

pictus, Anopheles (see A. hyrcanus).

Pigeons, distribution of Lynchia maura
on, 122.

Pigs, Oestrid infesting, in Africa, 247.

pilosa, Musca (see M. bezzit).

Pinnaspis buxi, food-plants of, in
Seychelles, 127.

minor, sp.n., food-plants of,
in Formosa, 214.

oe simplex, sp.n., in China, 214.

Pinus longifolia, new weevil bred from,
in India, 170.

thunbergii, Coccid on, in For-
mosa, 215.

pionips, Aédes.

pipiens, Culex.

Pipunculus annulifemur, sp.n., parasite
of Idiocerus in India, 465, 469.

Plasmodium spp., Leptoconops
experimentally infected with, 5.

Platyedrva gossypiella, on Cajanus indi-
cus in Mauritius, 182, 185.

Plecia fulvicollis, 232.

plinthopyga, Sarcophaga.

plumbeus, Anopheles.

Plusia (see Phytometra).

Plutella maculipennis, on cabbage in
Mauritius, 186.

Podocarpus chinensis, Coccid on, in
Formosa, 215.

Pollenia stygia, causing myiasis, 250 ;
larva of, 253.

pollinosa, Musca (see M. cingalatsina).

pomonae, Bibio.

pomonella, Cydia.

pomorum, Mytilaspis (see Lepidosaphes
ult).

Poppy, food-plant of Aphis rumicts, 81.

Porpe bjerkandrelia, on artichoke in
Mauritius, 186.

posticata, Psorophora.

Potassium Bitartrate, beetles apparent-
ly feeding on, 133.

Potato, attacked by Bibionid larvae,
232 ; food-plant of PAytometra ori-
chalcea, 99.

praecox, Musca (see M. humilis).

praematurus, Tabanus.

pratti, Cephenomyia.

primitiva, Musca (see M. humilis).

princeps, Bogeria.

priscus, Tabanus.

”

not

GENERAL INDEX.

Pristirhynchomyia lineata (see Musca).

Procervas sacchaviphaga, on maize in
Mauritius, 184.

Procontarinia matteiana, on mango in
Mauritius, 189.

Prodenia litura, food-plants
Mauritius, 181, 183, 184, 187.

promisca, Musca (see M. humilis).

Protomelanoconion, 79.

fuscum, referred to
subgenus Neocu-
Wee iS)

Psamma arenaria, new Aphid on, in
Britain, 430.

psammae, Laingia.

psarophanes, Silvius (see S. notatus).

Pseudaonidia aldabraca, sp. n., in

Seychelles, 125.

Oly Vin

”

- duplex, food-plant of, in
Formosa, 220.
- tola, sp. n., on Eugenia

caryophyllata in Sey-
chelles, 125.

* lacinia, 125.
a: tessevata, 126.
as trilobitiformis, food-plant

of; in Formosa, 220 ;
food-plant of, in Maur-
itius, 188, 189.
Pseudococcinella sexvittata (see Avrgo-
pistes).
Pseudococcus calceolariae, on ground-
nuts in Mauritius, 186.
i cityvt, food-plant of, in
Kormosa, 211 * on ‘cit=
rus in Mauritius, 188.

53 comstocki, food-plant of,
in Formosa, 211.
Me filamentosus, food-plant

of, in Formosa, 211;
on citrus in Mauritius,

188.

f vastator (see P. filamen-
tosus).

- virgatus, food-plant of, in

Formosa, 211.
Pseudogametes hermanni, infesting mice
in Brazil, 249.
re senratra, infesting mice
in Brazil, 249.
pseudonigeria, Aédes (Stegomyia).
pseudopictus, Anopheles (see A.
canus).
pseudopunctipennis, Anopheles.
Psorophora posticata, oviposition of, in
Trinidad, 481.
Pternoscirta minor, 162.
Piilolepis (see Musca).
Ptinus tectus, breeding in stored pro-
ducts) 134.
pugiunculus, Tabanus.
pugnax, Tabanus.

hyyr-

505

pulcherrimus, Anopheles (Myzomyia).
pulchripalpis, Orthopodomyia.
pulchritarsis, Aédes (Ochlerotatus).
pulicaris, Culicotdes.
pullatus, Aédes.
pumila, Musca.
pumilus, Culicordes.
Pumpkins, pests of, in
186.
punctibasis, Anopheles.
punctipes, Aédes (Skusea).
punctor, Aédes.
punctulatus, Anopheles.
punctum, Cratopus.
pungoana, Musca (see M. ventrosa).
purchast, Icerya.
puripennis, Culicoides.
purpurata, Rachionotomyia.
purpureus, Rhinoestrus.
pusillus, Culex (Barraudius).
Pycnosoma bezziana (see Chrysomyia).
Pygalataspis, gen. nov., 218.
rr miscantht, sp. n., on
Miscanthus sinensts in
Formosa, 218, 219.
pyophila, Sarcophaga.
Pyretophorus (see Anopheles).
Pyrilloxenos compactus, parasite of
Idiocerus in India, 465.
Pyroderces simplex, on
Mauritius, 182.

Mauritius,

cotton in

quadrimaculata, Calliphora.
quadrimaculatus, Anopheles.
quasigelidus, Culex.

quasimodestus, Culex (see C. pipiens).
quasiornata, Rachionotomyia(Stegomyia).
quatuordecimpunctata, Crioceris.

Rachionotomyia aenea, sp. n., in
Federated Malay

States, 79.
a bambusa, in Palae--

arctic Region, 284;
anatomy of, 284.
bimaculipes, 79, 80.
purpurata, sp. n., in
Fiji, 79, 80:
quastornata, in
tralia, 80.
m similis, 79.
Rachisoura filipes, synonymy of, 79.
os sylvestris (see R. filipes).
vadicicola, Heterodera.
Rape, food-plant of Phytometra ori-
chalcea, 99.
Red Scale (see Chrysomphalus aurantit) .
vegina, Phormia.
vegius, Megarhinus.
Reindeer, Oestrids infesting, 248, 250

Aus-

506

Remigia vepanda, on Vigna catjang in
Mauritius, 186.

vepanda, Remigia (Pelamia).

Rhadinomerus, compared with

cistocerus, 170.

‘3 bombacis, sp. n., in
Bombax malabaricum
in India, 170-172.

a diversipes, sp. n., bred
from forest trees in
India, 172-175.

- malloti, sp. n., on Mal-
lotus philippinensis
in India,175.

Ps subfasciatus, sp. n., in-
festing forest trees
in India, 176.

Rhadinopus buteae, sp. n., on Butea

frondosa in India, 177.

ihe centriniformis, 178.

consputus, 178.

i parcus, 178.

Rhinoceros, Oestrids infesting, 245.

Rhinoceros sumatrensis, Tabanid at-
tacking, in Malaya, 449.

Rhinoestrus hippopotamt, infesting hip-

popotamus in Africa,
247.

_ nivarleti, infesting pigs in
Africa, 247.

e phacochoeri, infesting wart
hog in Africa, 247.

y purpureus, causing myia-
sis, hosts and distribu-
tion of, 247.

Rhinotermes, 363.

Rhodesia, new Leptoconops in, 14.

vhodesiensis, Anopheles (Myzomyta) ;
Leptoconops.

Rhogas (Heterogamus) percurrens, Sp.n.,
parasite of Achaea janata in India,
131.

Rhopalocampta forestan, on Canavalia
ensiformis in Mauritius, 185.

rhynchias, Argyroploce.

vyichiavdii, Taeniorhynchus
tidia).

vodhaini, Covdylobia (Stasisia).

Roe-deer, Oestrid infesting, 250.

Rogenhofera spp., infesting mice in
South America, 249.

Rosa damascena, pests of, in Bulgaria,
353.

vosae, Aulacaspis.

Rose, Coccid on, in Formosa, 213 ;
attacked by Agvilus foveicollis in
Bulgaria, 353-354.

vostochiensis, Aédes (Ochlerotatus) cata-
phylla.

Rubber, scale-insects on, in Mauritius,
182.

vubens, Ceroplastes.

Me-

(Coquillet-

GENERAL INDEX.

vubicundulus, Tabanus.

vubidus, Tabanus.

vubriceps, Drepanotermes.

vubrolineata, Fiorinia.

vubus, Batocera.

vuficornis, Sarcophaga.

rufifacies, Chrysomyia (see C. albiceps).

vufinotatus, Tabanus.

vufipes, Aphiochaeta.

vufiventris, Cuterebra.

yufus, Culex (see C. pipiens).

Rumex, food-plant of Aphis rumicis, 87.

yumicis, Aphis.

Russia, breeding-places of locusts in,
155.

vusticus, Aédes (Ochlerotatus).

vutilus, Megarhinus.

sacchaviphaga, Proceras (Diatraea).

sacharovit, Anopheles.

sagitta, Gelastorrhinus.

Saissetia hemisphaerica, food-plant of,
in Formosa, 212; on citrus
and coffee in Kenya Colony,

103; food-plants of, in
Mauritius, 182, 184, 187,
188.

nigva, on citrus and coffee in
Kenya Colony, 103; food-
plants of, in Mauritius, 182,

187.
7 oleae, on citrus in Mauritius,
188.
salina, Culicada nemorosa (see Aédes
sticticus).

Saline Waters, mosquitos breeding in,
29-34.
salinellus, Aédes (Ochlerotatus).
Salix warburgi, new Coccid on, in
Formosa, 216.
salus, Culex (see C. sitrens).
sanguineus, Ividomyrmex.
Sapium sebiferwm, new Coccid on, in
Formosa, 216.
Sarcophaga, producing myiasis in India,
239; characters of lar-
vae of, 255, 256, 258,
260; larviparous habit
of, 261.
1 chrysostoma, causing myia-
sis, 250, 251.
oy lambens, causing myiasis in
South America, 251.
* plinthopyga, causing myia-
sis, 250; 251.
o pyophila, causing myiasis
in South America, 251.
a vuficornis, causing myiasis,
250.
savcophagoides, Cuterebra.
scalavis, Fannia.

GENERAL INDEX.

Scatopse, parasitised by Agyrtes bicolor,
231.
AF notata, 232.
Schistocerca peregrina, 159, note.
Schizoconops (see Leptoconops).
schmalzt, Cuterebra.
Schneideria mucronata, infesting Bibio-
nid larvae, 231.
scuddert, Bogeria.
scutellavis, Aédes (Stegomyia) (see A.
albopictus).

Selenaspidus articulatus, on citrus and
coffee in Kenya Colony, 103, 104.
selengensis, Anopheles (see A. maculi-

pennis).
semiatra, Pseudogametes.
semicantans, Aédes (Ochlerotatus).
separatus, Anopheles.
septentrionalis, Drepanotermes.
serenissima, Lucilia (see L. argyrice-
phala).
sergenti, Anopheles (Myzomyia) ; Culex.
sevicata, Lucilia.
Sesamia vuteria, bionomics of, on maize
in Mauritius, 183.
setigera, Musca (see M. albomaculata).
sexutttatus, Argopistes (Pseudococcinella).
seychellarum, Icerya.
Seychelles, Coccidae in, 125-128.
Shanghai, breeding-places of mosquitos
at, 403.
Sheep, Oestrids infesting, 244, 246.
Shorea robusta, new weevils bred from,
in India, 175, 177.
Siam, new Leptoconops in, 20; Taba-
nids of, 431-455.
stamensis, Leptoconops ; Tabanus.
Siberia, new mosquito in, 308.
stberiensis, Culiseta (see Theobaldia
alaskaensis).
sicavius, Tetrastichus.
Sida, Chionaspis subcorticalis on, in
Seychelles, 127.
silenus, Hypoderma.
silvestvi1, Diversinervus ;
mes.
Silvius fuliginosus (see S. notatus).
,» notatus, 55; bionomics of, in
Australia, 59-62.
,, psarophanes (see S. notatus).
similis, Haematopota; Microplitis ;
Rachionotomyia.
simplex, Pinnaspis ;. Pyroderces (Ana-
tvachyntis).
simpsoni, Culex.
Simulium, 1.
=i alcockt, sp. n., in Nigeria,
459.
Se alcockt var. coalitum, n., in
Nigeria, 460.
a alcocki var. violaceum, N.,
in Nigeria, 460.

Drepanoter-

507
Simulium aureosimile, 461.
‘ beckeri, in Algeria, 117.
. divergens, sp. n., in Nigeria,
460.
Ws equinum, attacking horses in
Palestine, 117.
- flavipes, sp. n., in Palestine,
116.
Re hivsutum, sp. n., in East
Africa, 458, 460.
- hivsutum var. aderysi, n., in

Zanzibar, 459.

a hivsutum var. dubium, n., in
East Africa, 459.

oy lineatum (see S. maculatum).

Fe maculatum, 4.

- neavet, 461 ; possibly carry-
ing human disease in
Kenya Colony, 236-238.

e palmeri, sp. n., in Nigeria,

462.
ue unicornutum, in Nigeria, 462.
rf vorax, sp.n., in East Africa,
461.

sinensis, Anopheles (see A. hyrcanus) ;
Culex ; Mallococcus.

sinicus, Tabanus.

Sipha, Laingia allied to, 429.

sitiens, Culex.

smitht, Lachnosterna (Phytalus).

solani, Coelonia (Sphinx).

sordidus, Cosmopolites.

spathipalpis, Culex
longiareolata).

spectabilis, Microplitis.

speculifera, Musca.

Sphinx convolvuli (see Herse).

,, solani (see Coelonia).

spinohumera, Musca.

spinosa, Musca.

spinosifrons, Acanthoconops.

spinosipes, Leptoconops (see Acantho-
conops albiventris).

Spodoptera mauritia, food-plants of, in
Mauritius, 181, 183.

Stasisia vodhaint (see Cordylobia).
Stauropodoctonus mauritii, parasite of
Sesamia vuteria in Mauritius, 184.
Stegomyia, key to Palaearctic species of,

324 ; asubgenus of A édes,
293, 324 5 G-0.
- hilli (see Rachisoura filipes).

(see Theobaldia

ie quastornata (see Rachiono-
tomyia).
55 microptera, 77.

- periskeleta, 77.
stellifera, Vinsonia.
stephenst, Anopheles (Myzomyia).
sticticus, Aédes (Ochlerotatus).
stimulator, Cephenomyia.
Stomoxys, possibly transmitting try-
panosomiasis to cattle, 236.

508 GENERAL INDEX.

Stomoxys calcitrans, attacking cattle in
Kenya Colony, 235; in
Palestine, 121.

nigva, attacking cattle in
Kenya Colony, 235.

varipes, attacking cattle in
Kenya Colony, 235.

striatecta, Musca.

stviatus, Cevatosolen ; Tabanus.

stvgia, Pollenia.

stygius, Leptoconops.

subcorticalis, Chionaspis.

subfasciata, Rhadinomerus.

subochrea, Theobaldia.

Sudan, new mosquito in, 69.

Sugar-cane, Aulacaspis tegalensis on, in
Formosa, 213.

sugens, Aédes
vittatus).

sulcicollis, Locusta (see Locustana par-
dalina).

sumatrensis, Gyrostigma.

Sunflower, food-plant of <A sterochiton
vaporariorum in Britain, 359.

superpictus, Anopheles (Myzomyia).

suvcoufl, Kirkioestrus.

susae, Culicoides.

Sweet Potato (Ipomoea batatas), pests
of, in Mauritius, 184.

sygmotdes, Dacus.

syluae, Culex nigripes (see Aédes sticti-
cus).

sylvestris, Rachisoura (see R. filipes).

Sympiezomias beesoni, sp. n., on teak in

India, 165.
- frater, 166.

”

(Stegomyia) (see A.

Tabanidae, bionomics of, in Australia,
41-62; new species of, in Siam,
431-455.

Tabanus, key to Siamese species of,
433-435.

albimedius (see T. priscus).

agnoscibilis, sp. n., in Siam,

453.

Fe apvepes, bionomics of, in
Australia, 41-53.

A barnesi, sp. n., in Siam, 435-
ASTe

s batchelori (see T. aprepes).

a brunnipennis, in Siam, 444.

a cinerascens, in Java, 453.

oe designatus (see T. rufino-
tatus).

7" elestéem (see T. rufinotatus).

~ finalis, in Siam, 442.

im fulvimedioides, in Formosa,
451.

ie fulvimedius, in Formosa, 451.

oh hilaris (see T. tenens).

a indianus, distribution of, 444.

Tabanus insidiatoy, sp. n., in Siam,
437-440.
= (A tylotus) leucocnematus, 439—
440.
FF lineatus (see T. vufinotatus).
as megalops (see T. tenens).
¥ (Atylotus) nephodes, in Assam,

437.

ae nigvitarsis, 43 ; bionomics of,
in Australia, 56-59.

#3 nigrotectus, in Siam, 437.

ie partitus (see T. striatus).

praematurus, sp. N., in Siam,
440-442.

a priscus, 448.
> pugiunculus, sp. n., in Siam,

451453.

is pugnax, sp. n., in Siam,
449-451.

Be yvubicundulus, sp. n., in Siam,
442-444.

e vubidus, in Siam, 448.

vufinotatus, 41, 43 ; bionomics
of, in Australia, 53-56.

siamensis, in Siam, 455.

He sinicus, distinct from T. stv1-
atus, 446.
- stviatus, distribution of, in

Far East, 445.
taeniola var. variatus, 235.

* tenens, 445.
5 virgulatus, sp. n., in Siam,
446-448.

Tachardia decorella, food-plant of, in
Formosa, 212.

Tachina mella, 4.

Taeniorhynchus (Coquillettidia) vichi-

ardit, in Palaearctic Region, 291.

Taeniorhynchus titillans, recorded in
error from Rumania, 291, note.

taentorhynchus, Culex.

tahoensis, Aédes.

Tanganyika Territory, new Simuliids
from, 458, 461.

tavandi, Oedemagena.

tavandus, Oryctes.

tarvsimaculatus, Anopheles.
Tarucus telicanus, on Cajanus indicus
in Mauritius, 185.
tasmaniensis, Aédes
Lucilia.

taylori, Culex.

Teak, new weevil attacking, in India,
166.

tectus, Ptinus.

tegalensis, Aulacaspis.

Telenomus, parasite of Prodenia lituva
in Mauritius, 181, 184.
truncativentris, parasite of
Antestia lineaticollis in
Kenya Colony, 191-201.

telicanus, Tarucus.

(Ochlerotatus) ;

GENERAL INDEX.

tempestiva, Musca.

tenax, Evistalis.

tenebrosa, Cutervebra.

tenens, Tabanus.

tentorius, Culicoides.

Termites, new species of, in Australia,
363-399.

ternicinctus, Gastrophilus.

tervae-veginae, Musca.

Tersesthes, a synonym of Leptoconops,
£2 107,

tessellatus, Anopheles
Eucalymnatus.

tessevata, Pseudaonidia.

Tetrastichus sicarius, parasite of Coccus
viridis in Mauritius, 188.

thalassius, Culex.

Thaumasura curculionis, parasite of
Euthyrrhinus meditabundus in
Queensland, 65.

Thea sinensis, Coccid on, in Formosa,
214.

theobaldi, Anopheles.

Theobaldia, key to Palaearctic species
of, 286.

alaskaensis, in Palae-
arctic Region, 288.

annulata, in Britain, 208,
427; in Palaearctic
Region, 288; anatomy
of, 266, 292.

arctica (see T. alaskaensis).

bergrotht (see T. glaphy-
voptera).

dyari, 289.

(Culicella) fumipennis, in-
fluence of drought on,
in British Isles, 207; in
Palaearctic Region, 290 ;
anatomy of, 292.

punctulatus ;

- (Theobaldia) glaphyvop-
teva, in Palaearctic Re-
gion, 287.

(Allotheobaldia) longi-
aveolata, in Palaearctic
Region, 287; venation
of, 285.

(Culicella) morsitans, in-
fluence of drought on,
in British Isles, 205,
207; in Palaearctic
Region, 289.

(Pseudotheobaldia) nivei-
taeniata, distinct from
T. glaphyroptera, 287.

(Theobaldia) subochrea, in
Palaearctic Region, 289.

Theretva alecto, bionomics of, on grape
vines in Mesopotamia, 478.

Thliptoceras octoguttalis, Botys octo-
guttatus possibly recorded as, on
coffee in Mauritius, 187.

”

509

thoracicus, Culex.

Thrips, food-plants of, in Mauritius,
183, 185.

Thripsaphis, Laingia resembling, 429.

tibeviadis, Trichotanypus.

tigvipes, Lutzia.

tipuliformis, Culex.

titillans, Taeniorhynchus.

titillator, Cephalopsis.

Tobacco, food-plant of Asterochiton
vaporariovrum in Britain, 359 ; pests
of, in Mauritius, 182.

togot, Aédes (Finlaya).

Tomato, Astevochiton vaporariorum on,
in Britain, 355-359; pests of, in
Mauritius, 186; Chionaspis sub-
corticalis on, in Seychelles, 127.

torrens, Leptoconops.

towadensis, Megarhinus.

Toxoplasma, 5.

Transcaspia, breeding-places of locusts
in, 143-154.

Trichogramma australicum, parasite of
Sesamia vuteria in Mauritius, 184.

Trichonympha, in termites, 371.

Trichoptilus wahlbergi, on sweet potato
in Mauritius, 184.

Trichotanypus tiberiadis, recorded in
error as a blood-sucker in Palestine,
ELS;

trifilatus, Culex.

trigonocephala, Rogenhofera.

Trigonogenius globulum, breeding in
argol, 133.

trilobitiformis, Pseudaonidia.

Trinidad, oviposition of Psorophora
posticata in, 481.

Trioza, on citrus in Kenya Colony, 103 ;
on citrus in Mauritius, 188.

tritaentorvhynchus, Culex.

tvompe, Cephenomyia.

Trullifiorinia, 216.

truncativentris, Telenomus.

Trypanosoma uniforme, in cattle in
Kenya Colony, 233.

Trypanosomiasis, in the absence of
Glossina in Kenya Colony, 233-236.

tubulorum, Lepidosaphes.

tuvanica, Oxya.

turkhudi, Anopheles.

tyriet, Eutermes.

ugandae, Haematopota.
ulmi, Lepidosaphes.
ulrichi, Cephenomyia.
umbrosus, Anopheles.
unguiculata, Uvanotaenta.
unicornutum, Simulium.
uniforme, Trypanosoma.
univittatus, Culex.

510

Uvanotaenia bimaculata, in
arctic Region, 283.
mashonaensis, 283.
ae unguiculata, in Palae-
arctic Region, 283.

vagus, Anopheles.

validicornis, Haematopota
pachycera).

validus, Leucotermes.

vaporarviovrum, Asterochiton (Aleurodes).

variatus, Tabanus taeniola.

variegatum, Amblyomma.

varioannulatus, Culex (see C. pipiens).

variolus, Oestrus.

varipes, Chrysomyia ; Stomoxys.

vassilievi, Anopheles superpictus.

vastator, Pseudococcus (see P. filamen-
tosus).

vastatrix, Hemileia.

Vegetable Marrow, food-plant of A stevo-
chiton vaporariorum in Britain, 359.

venosus, Bibio.

ventyalis, Armigeres.

ventrillont, Culex.

ventrosa, Musca.

Verschaffeltia splendida, new scale on,
in Seychelles, 127.

versutus, Adoretus.

verticillata, Eupristina.

vetevinus, Gastrvophilus.

vetustissima, Musca (see M. pumila).

vexans, Aédes (Ecculex).

Viburnum opulus, food-plant of Aphis
vumicis, 86, 87.

vicavia, Musca (see M. domestica).

victoriensis, Hamitermes perplexus.

Vigna catjang, pests of, in Mauritius,
186.

Vine, Grape, Lepidopterous pests of,
in Mesopotamia, 478, 479.

Vinsonia stellifera, on mango in Mauri-
tius, 189.

violaceum, Simulium alcockt.

virgatipes, Culex.

virgatus, Pseudococcus.

virgulatus, Tabanus.

(see A.

Palae- |

GENERAL INDEX.

viridis, Agrilus ; Coccus; Culex.
viridissima, Crioceris.
viridula, Cochliomyia.
vishnut, Culex.
vitreipennis, Culicoides.
vitvipennis, Musca.
vittatus, Aédes (Stegomyia).
vivax, Plasmodium.
vomitoria, Calliphora.
vorvax, Lutzia; Simulium.
vulgaris, Culex.

vuleria, Sesamia.

wahlbergi, Trichoptilus.
Wart Hog, Oestrid infesting, in Africa,
247.
waterhousei, Aédes (Ochlerotatus) (see
A. maculatus).
Weevils, new, in India, 165—180.
willmori, Anopheles (see A. maculatus).
Willow, new Coccid on, in Japan, 216.
Wohlfahrtia, larviparous habit of, 26M.
- magnifica, causing myiasis,
241, 243; larva of, 260.

xanthina, A phiochaeta.

xanthindyma, Cosmophila (see C. flava).

Xanthogramma pfeifferi, predacious on
Aphis brassicae in Mauritius, 186.

xanthomela, Musca.

Yokohama, breeding-places of mos-
quitos at, 407.
ypsilon, Agrotis.

zammittit, Aédes (Ochlerotatus).

Zanzibar, new Leploconops in, 24; new
Simulium in, 459.

Zebra, Oestrids infesting, 245.

zebrae, Gastrvophilus pecorum.

Zizera lysimon, on Cajanus indicus in
Mauritius, 185.

zizyphi, Parlatoria.

511

INDEX TO NAMES OF PERSONS.

Abildgaard, Prof. P. C., 416.

Adams, J. J., 103.

Adelung, Dr. N., 141, 288.

Aders, Dr. W. M., 26, 242.

Ahnper, C:,..273, 275; 279, 287,. 288,
300, 323, 336.

Alcock, 1t.-Col. A. 79),.94, 120; 281.

Aldvich, OT. [iM 24s 05,717, 23

Alluaud, 316.

Anderson, T. J., 99, 191, 195, 233, 473.
Annandale, Dr. N., 20, 115, 119, 120,
note, 121, 287, 279, 345, 432, 453.

Artsimovicz, V., 135.
ASaworth, Dr). EL. 76.
Aurivillius, Prof. C., 306.

Austen, Major E. E., 107-124, 263,
IN 33d, (309, O42, o40, 411, 412,
413. 415, 416, 417; 418,’ 420; 421.

431-455.
Averill, Capt. W. W..,.122.
Awati, 416, 418, 420, 421, 422.

Bacot, A., 74:

Bahr 2Dr, 2 El wor

Baini Prashad, Dr., 423.

Balfour, Dr. A., 29-34.

Bancrort, Ors i. i fo; 425;

Banks, 250, 254; 284° 329) 327%

Banber, (os. lde io.

Barnes, Dr. M. E., 431, 432, 437, 441,
442, 449, 451, 453, 455.

Barraud, Capt...P. .J.,, t16,. 263; 27.1,
DUS OTROS LOU 280) 300303;
SUA WaZay S20; S00; O04, O09, O40;
342, 343, 344, 345.

Barrow, 328.

Bartko, 300, 302.

Bate, Miss, 338.

Baume Wr la TS) 152.

Bedford, 340.

L762 ITS als:
Bell-Marley, H., 475.

Bengtsson, Dr., 320.

Bennett, Major, 77.

Bequaert, Dr. J., 240, 245, 246.

Bergroth, Dr. E., 263, 287, 311, 314.

Berland, 304.

Bezzi, Prof. M., 3, 14, 17, 263, 291,
304, 316, 323, 324, 342, 420, 422,
423.

Bigot, 326, 333, 413, 418,
433, 437, 440.

Bird, 300, 302, 306.

Bischoff, 307.

Bishopp, F. C., 240, 241, 251.

Blane, ‘G.,. 5.

Boag, G., 342.

Boden Kloss, C., 7

Boheman, C. H., 2
S10) 314, S16, 320;

Boldyrev, V., 136.

Bouet, 242.

Brain, Wr. C. K., 475: .

iBrandeckt so. p409)1"

Brauer, 244, 288, 418, 422.

Bredemann, 479.

Brolemann, H. W., 286, 312, 316.

Brunetti, E., 115, 121, 469.

Brunner v. Wattenwyl, 152.

Bryant, G. E., 473-475.

Butler, Dr. G., 74.

Butler, Capt., 437.

Buxton, P. A., 323, 341, 477.

419,

IQs
88, 291, 304, 305, 307,
20.320:

Cameron, Capt. J., 233, 234.
Carls Dry. j., 162:

Carment, Capt., 309.
Carpenter,.Dr. G.H., 232, 248.
Garry olde

Carruthers, 30.

Carter Heb 1-28, 107, 33:
Castellani, A., 1, note, 239.
Chalmers, A. J., 1, note, 239.
Champion, G. C., 134.
Champion, H. G., 170.

Ee, INDEX TO NAMES OF PERSONS.

Chapin, C. V., 30.

Chari, 248.

Charmoy, D. d’Emmerez de, 181-190.

Chatterjee, N. C., 166

CGhattons E375:

Cheah, Dr., 409.

Chidester, F. E., 31.

Chopard, L., 163, note.

Christophers, Major S. R., 29° 303, 267;
270 27 272) Zio Oe es
279, 280, 327, 428.

Chuen, M. C., 97.

Clank, 5,007.

Clearkin, Dr., 238.

Cobbet, L., 29.

Cockayne, Dr. E. A., 309.

Colclough, S., 473.

Coleman, Dr., 467.

Collet Prot. Ox owe

Collin, J. E., 320, 333, 413, 417.

Corfield, Major W. F., 119.

Cornford, 327.

Cragg, Major F. W., 92, 120, 412, 415,
420, 421.

Cropper, 338, 343.

Cross, Capt., 248, 261.

Csiki, 308, 313.

Daking jew; 12.

Dale, Capt. W. J., 121, 122.

Dalziel Dr J. NM. 29, 30, 31:

Darling. S- T.; 3:

Davidson, J., 81-89.

Davies Sherborn, C., 267.

Davis er iC. Ne 4092

d’Emmerez de Charmoy, D., 181-190.

de Fourcroy, 288.

de Graaf, J. M. H. Swellengrebel, 70.

de Lezey, L. Drouard, 284, 285, 318,
O27.

de Meijere, Prof. J. C. H., 2,3, 26, 78,
304.

de Vogel, W. T., 30.

Debreuil, 313.

Dobbs. .€. ME, 235:

Dodd, A. P., 65, 66, 67-68, 191.

Donitz, 70, 71, 280.

Dootson, Dr. F. W., 133.

Drake-Brockman, R. E., 31.

Drouard de Lezey, L., 284, 285, 318, 327.

Dry, F. W., 99-102, 103-104, 191-201,
233-238.

Duffield, Capt. A., 430.

Dupont, Pi R., 125, 126, 127.

Durrant, J- H.,-137, note:

Dict. eles, N29 SZ.

Dutton, J: 2.7729:

Dyar, Hi: G:; 30) 31,192,267, 280;-28L,
284, 288, 289, 292, 293, 301, 309,
SOs Sid) Sl6, 322) SZS S262,
SISMSSOUS46No47e

Eaton, Rev. A. E., 122, 319:

Eckstein, F., 288, 312, 313, 323, 336.

Edwards, F. W., 2, 69-80, 116, note,
117, 221, 236, 263-351, 406, 407,
409.

Enderlein, Dri (Gs) 23) 78, 263.92775
302, 306 mol7e

Essen, Wav, G06) oN ole:

Essig, Prof, H20., 241;

Evans, A. E., 474.

Falshaw, P. S., 449.

Faure, J. C., 136, 155, 157, 158, 159,
475.

Ferguson, Dr. E. W., 419, 423.

Ferris, G. F., 211-220.

Ficalbi, E., 263, 287, 290, 304, 305,
307, S12" 333; Goo, O47.

Field, J. F., 378.

Fielding, J. W., 31.

Finsch, 314.

letchers i. Bey 129:

Flower, Major S. S., 445.

Foley, H., 30, 280.

Bord, C.7.302%

Forsius, 311.

Forskal, 347.

Fortunat, 327.

Fowler, 321, 344.

Frauenfeld, 291.

Frey, Dr. R., 263, 287, 306, 311, 314,
BLOF

Froggatt, W. W., 251, 376, 419, 423,
424,

Buller G27 363:

Gahan, DriC. J., 178:
Gairdner, K. G., 442, 451, 454.
Galli-Valerio, B., 316,319)
Gaillois, E:, 318; 327,339.
Gebert, S., 181-190.

Gedoelst, Dr., 244.

Gill, 342, 344.

Gjellemp, K., 26.

Glaser, 248.

Gliniuk, T., 136, 143, note.
Godfrey, F. J., 445, 448.
Goetghebuer, M., 304.
Goldberg, Dr., 289, 300, 339, 342.
Gough, Dr. U5 Hy 4,30:

Graaf, J. M. H., Swellengrebel de, 70.
Graham, W. M., 29.
Graham-Smith, 342.

Gran6, 308, 324.

Grassis Bin 0n Zo eior
Gravely, F. H., 73.

Green, E. E., 125-128, 211, 214.
Griinberg, 303.

Giinther, 288.

Se oe one

INDEX TO NAMES OF PERSONS.

Hacker, Dr. H. P., 70, 91, 406.

Hadwen, Dr., 248, 249.

Haglund, 288, 302, 304, 305.

Handlirsch, A., 288, 300, 302, 313, 316,
336.

Hargreaves, E., 263, 271, 283, 304, 308.

Harmand, 318.

Hartert, Dr ES 107:

Harvey Johnston, Prof. T., 423, 425.

Heller, Prof. K., 170.

Hewitt. Dr: C2 G4 11 412:

Hewitt, J.;35, 38; 40.

Hill, G. F., 41-62, 63-66, 71. 75, 76,
78, 79, 80, 363-399, 373, 376, 419,
423, 424, 425.

Horn, Dr. W., 263.

Horvath, Dr. G., 304.

foward, “Dr L7@:; 2) 15, 23, 30931;
924280). 281, 284.301, 323, 328;

Hubback, T. R., 449.

Hudson,G. Vv, 73: 74) mote:

Ikonnikov, N., 136.
Imms; Dr: A.D. 2271:
Inamura, S.. 211:
Ingram, Dr. A., 74, 342.

Jack; R. .L.,. 386.

Jack, R. W., 242.

Wacoby jE. S2.

james, <1t:-Col. Sv P4279, 427-499;
Jatzentkovsky, E., 135.

Johannsen, O. A., 1, 2.

Johnston, Prot:S- 7... 10,

Johnston, Prof. T. Harvey, 423, 425.
Joyeux, C.. 242,275, 283.

Karsch, F., 341, 419.

Kearmey, W.,.233:

Isetline De. 23h 239%

Kellogg/V.R., 211.

IMCD, .9., 7 2:

Kendle, Capt. 117.

Kertesz,. Dr. 1¢., 263, 288))-291>- 300:
302, 306, 308, 311, 316, 333, 433
note.

Kieffer, J jij) note, 2; 3;.18;-19, oR
D3 NOG:

Kilner, E., 404, 409.

King, H. H., 29, 69.

Kong, We J. E341.

Kirby, W. F., 137, 162.

Kloss, C. Boden, 72.

Knab, F., 30, 31, 92, 267, 280,
284, 301, 323, 327, 328, 346.

Koidzumi, M., 319.

Kozhevnikov, Prof. J., 135.

Krauss, H., 136.

Krogerus, 288.

281;

513

Kuntze, 304.
Kuwana, 211, 214, 217, 219.

La Baume, Dr., 151, 152.

Labbe, P., 306.

Laing, F., 125-128, 429.

Lamborn, Dr. W. A., 91-97, 263, 274,
318, 325, 327, 328, 337, 339, 342,
345, 401-409.

Landrieu, Dr., 335, 336.

Lang, Dr. W. D., 273, 291, 304, 323.

Langeron, Dr. M., 3, 4, 5, 6, note,
22, 279, 280, 287, 288, 300, 303, 304,
336, 342.

Lea, A. M., 391.

Leicester, Dr. G. F., 70, 79, 283.

Lelean? Lt.-Col.; 272.

Lendl, 300.

Leon, N., 275, 291, note, 333.

Leonardi, 216.

Lesne, P., 312, 313.

Levander, 314, 323.

Lezey, L. Drouard de, 284, 285, 318,
327.

Lichtwardt, B., 263, 290, 300, 302,
304, 306, 311, 313, 314, 316, 325.

Liston, 279.

Lloyd, Dr. Ll., 355-359.

Lyle, G. T., 129-132.

Loew, H., 317.

Loitsch, 300.

Ludlow, Dr. C. S., 288, 318.

Lundbeck, Dr., 263, 309.

Lundstrém, 288, 290, 300, 302, 307,
314.

Lutz, Dr. A... 3,135.

Macdonald, 280.

Macfarlane, 342.

Macfie, Dr. J. W. S., 31, 342.

MacGregor, M. E., 29, 205-209, 291,

427, 428.

Macleod, Capt. E. C., 440.

Maki, M., 211.

Malaquin, 87.

Mann, 300, 305, 307, 308, 311, 313,
314, 320, 336.

Marshall, Dr. G. A. K., 2, 153, 165-180,
240, 261, 268, 414, 426.

Martini, Dr. E., 263, 300, 305, 307, 311,

315, 316; 323, 338:

McDonald, J., 100.

Meijeres Prot: Je'C. H. de. 2 37036.

78, 304.

Metcalf, 253.

Mik, 1, 288, 302, 305, 308, 313, 314,
S16; 3335-8386.

Miller, D., 74, note.

Mills, Dr. R. G., 318.

Mitzmain, M. B., 445, note.

514

Mj6berg, Dr: E., 363, 379,378; 382,
386.

Moitié, 87.

Mokrzecki, Prof. S. A., 353-354.
Molz, 5 .,.232-

Montgomery, R. E., 233.
Mordwilko, A., 87.

Morgan, J. de, 289.

Montz. 1s. 036:

Morris, Ee VE, 221=232:
Motchulsky, 307.

Mouhot, 437, 444.

Naday, 308, 323, 333, 341.

Neave, Dr. S. A., 473.

Neveu-Lemaire, 285.

Newstead, Prof. R., 113, 213.

Noe, G., 2; 3, 4, 5; 6, note, 14508. 338r
Nuttall, rot G. EE 29273.

Oldenberg, L., 263, 273, 304, 305, 306,
SOM molars lAataoos

Orchardson, I. Q., 233, 235.

Osmaston, B. B., 180.

Pachossky,, J. K., 135.

Palmen, 316.

Palmer, W., 432.

Parker, 249.

Patterson, W. H., 474.

Patton, Major W. S., 92, 120, 239-261,

411-426.

Pavie, 432.

Pawan, J. L., 481.

Pendlebury, W. J., 289.

Penther, 316.

Perkins, M. G. L., 134.

Rerkins Dre Cole A468:

Perrone, E., 30.

Perry; J. €., 32:

Peryassi, A. G., 31.

Pettigrew, Rev. W., 442.

Pic, M., 134.

Pipping, 314.

Pirkovsky, G., 136.

Plotnikov, V. J., 135, 148, 149, 150.

Pokorny, 288, 307, 311, 313.

Pomeroy, A. W. J., 457-463.

Portchimsky, Prot. 1yA2, 243; 285, 42):

Prashad, Dr. Baini, 423.

erartaty Euan Coe elie

Pratt, H. C., 135, note, 136, 142, note,
NS ta:

Priestly, Dr. 424.

Racquette, 310.
Ramachandra Rao, R., 477-479.
Rebn, Dr. J:, 136, 152; note,

INDEX TO NAMES OF PERSONS.

Ricardo, Miss G., 432, 445, 446, 455.

Richardson, Lt.-Col., 118.

Riedel, 304, 305, 311.

Robinson, H.C., 20, 432.

Rodhain, J., 240, 245, 246.

Roepke, Dr., 153,

Roman, Dr., 420.

Roper, Dr. 70:

Rossikov, K. N., 146.

Rothschild, IN. C., 412.

Roubaud, Dr. E., 240, 242, 243, 244,
278.

Rovere, 242.

Rutherford, A., 128.

Sahlberg, J., 290, 302, 306, 311, 314,
316.

Sahlberg, U., 314.

Sakharove Nodes oo:

Sambon, Drei. We; 40:

Sander Drs les Lolo:

Sangone, 304.

Saunders, W. W., 437, 444.

Schiner, 287, 288, 304.

Schneider, P., 313, 323.

Schonherr, 288.

Schulz, 302.

Scott. Dre Ee 133-134"

pearle, CaptsC., 121°

Séguy, E., 263,285, 291, 301, 303, 304,
306,312, 315; 316, 320, 336.7340)
note, 342.

Sella, M., 31.

Semenov-Tjan-Shansky, A. P., 153.

Senior-White, R., 415, 419, 421, 422.

Sergent, E., 270,271, 279) 2800303;
oeile

Sewellmlet.-Colnea ese 0a le

Sharpe Drs loo mote:

Sherborn, C. Davies, 267.

shevyrev, 1.) 137; note:

Shipley, Sina E273.

Slovene, lel, 123, Avex.

Shtchelkanovzev, Prof. J., 135.

Shute, P. G., 427.

Silén, F., 314.

Silvestri, Prof. F., 398.

simpson, Dr Je jc, lor.

Sj6stedt, Prof. Y., 162, 263, 309.

Skuse, Av Ale 7.18

Solier, 133, note.

Somina, Miss. O. M., 137, note.

Sorauer, P., 231.

Stanton, Dr. A. T., 401, 408, 432.

Stein, Prof., 418, 419.

Stobbe, 305, 306, 314.

Storey, G., 334, 342, 343, 345.

Strangeways-Pigg, T., 29.

Strickland, 326.

Strobl, 316.

Subramaniam, T. V., 465—467, 468, 469.

INDEX TO NAMES OF PERSONS.

Summers, Miss S. L. M., 120.

Surcouf, Baron, J. M. R., 432, 437.

swellengrebel, Dr. N. H., 30, 70, 71,
WOre27 or .

Swellengrebel de Graaf, J. M. H., 70.

Tams, W. H. T., 468.

avlor yk’: Male 3 lee let Het LOM OO, 419:
423.

Maylor, i. e457,

Tetens, 307.

Thalhammer, 302.

Theobald, F. V., 87, 287, 300, 303, 304,
305% ole OS; 2 olON 45005, oo7., O39)
346, 347, 408, 429-430.

Thomson, 418, 420, 425.

Tindale; N. B..391;

Tonnoir, A., 305.

MRownsend GC mE Tol A bee 15< 16:
Moves Wa LOT, lal 2454 249,
irybom, 288, 306), 308.311, 313, 314:

324.

Uhl, 300.

Ujhelyi, 300, 302, 304, 306, 308, 311,
314.

Uvarov, B: P:, 135-163.

Vassitliev,, J. By 277.
Velitchovsky, 302, 311.
Villeneuve, Dr. J.,:287;:316; 420.
Vinokurov, G., 136.

Vivante, 30.

Vogel, W. T. de, 30.

von Bodemeyer, 336.

von Wattenwyl, Brunner, 152.

515

Wahlberg, P., 306, 314.
Wallace, A. R., 418.
Wanen, 307.

Waterston, Dr. J., 35-40,
213}; 2795" 2830 285,289,
CWA Ly sroto top aroretey.

Watson, Dr. M., 91.

Weiss; A.) 3,4, 14. 15> 21. 29—93)

Wells, 249.

Wenyon, Dr. C. M., 124, 273.

Wesenberg-Lund, Dr. C., 263, 272, 289,
293, note, 300, 301, 302, 303, 304,
306,/307;-310, S11, 313,.316).347.

Wesselman, H., 375.

White, S. A., 378.

Wildermuth, C. K., 4.

Wallcocks, F.C. 4. 21) 99:30:

Williamson, Dr. G. A., 273.

Wilson, F. E., 389.

Winthem, 305.

Wood, C. H. B., 408.

Woods. 15:

Woolley, Dr. P. G., 482, 445, 453.

Wuorentaus, 288, 308, 323, 342.

Wyville-Thompson, 342.

263,
304,

2715
308,

Yamada, S., 263, 283, 284, 318, 319,
GLO O20) ODF Oo, Goo:

Yerbury, Col., 320:

Yvernault, A., 30.

Zehntner, 213.
Zemey,; IT. 1. 260:
Zetterstedt, 301, 305, 307, 320.

weil &
os TAL =P, of
at As ft ehetviiles

Tire a! (ni, , CA
A ee PL) |

Me ai - rs
het i res 0 i) ey ae pas
fp wey OR ae ie. tay) ete
iL Ete TEEN ned

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L’ AGRICOLTURA COLONIALE (FLORENCE) :

Anno XIII (1919). No. 2.
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Voll -11(1902) Nos) 3 anda viol. lV (L905) seNor/S

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AMERICAN NATURALIST (NEW YORK):
Nos. 309, 315 to 317, 328, 369, 402, 441, 495, 497 to 500, 511; indices to,
Vols. XXVII and XXX; and pp. 423-438 of Vol. XXXII.
ANNOTATIONES ZOOLOGICAE JAPONENSIS (TOKYO) :
Vols. I-II (1897-98). Entire.
Vol. III (1899). No. 1.
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Vols. I-II (1906-10). Entire.
Vol. TW (1911=12). “Now 1:
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Reports on Insects and Fungi Injurious to Crops (Proceedings under the:
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BULLETIN OF THE ILLINOIS STATE LABORATORY OF NATURAL HISTORY (URBANA) :
Vol. VII (1904-12). Articles III to IX; XI to end and Index.
Vols. VIII and IX. Entire.
BULLETIN DE LA SOCIETE PORTUGAISE DES SCIENCES NATURELLES (LISBON) :
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Vols. I-IV (1889-94). Entire.
GEORGIA STATE BOARD OF ENTOMOLOGY (ATLANTA) :
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Girculars: 1 to 6,12, lo tol'8;and)20:
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SI Raho EE st EL

ie The Editor will be pleased to receive for publication papers or
_ notes dealing with any insects that are of economic
Such communications to be addressed to

importance.
THE DIRECTOR,
Imperial Bureau of Entomology,
British Museum (Natural History),
London, S.W.7,

The publication and distribution of the ‘‘ Bulletin of Entomological
‘Research ”” has now been taken over by

THE ASSISTANT DIRECTOR,
Imperial Bureau of Entomology,

| | : 41, Queen’s Gate, London, S.W.7,

ve.
i

| and orders and subscriptions should be sent direct to him or eae
| any bookseller.

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(2416)

CONTENTS.

ORIGINAL ARTICLES.

BALFOUR, Dr, ANDREW. ‘Miser Breeding in eat Waters. :

CarTER, Henry F. A Revision of the Genus Leptoconops, o

Skuse (illustrated) .. S ee ae ie sc ue

Davipson, Dr. J. Piles Studies of Aphis rumicis, oa
Linn. (tilustrated) 5

Dopp, ALAN P. A New Chalcid Parasite of bathyrhin
meditabundus .

Dry, F.W.. Flax aa in Kenya Colony, with ee oe
Reference to the Limitations of the oe Method of —

Combating Them .. oe
Dry, F. W. The Red Scale, Chomp aurantii, Mask, |
in Kenya Colony = .. s : i ee

Epwarps, F. W. Mosquito Notes,—lIl.

Hitt, G. F. . The Bionomics of Tabanus aprepes, and other
Australian Tabanidae (2/lustvated) . ee

Hitt, G. F. The Life-history of eee ee :
Fabr., an Important Weevil Pest of Mango Trees in
Australia (tlustrated) - es os 3 BAS Ne

Lamporn, W. A. The Nature and Function of the baadal Ma
Tufts of Malayan Anopheline Larvae (illustrated) —-

WATERSTON, JAMES. On some Bornean bogs
(Agaonidae—Hymenoptera Chalcidoidea) pao

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fs; : ; Pa

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a a J

A a

The: ‘Editor will be’ pleased ‘to receive for publication papers or
“i notes. dealing with any Insects that are of economic importance.
‘ Such communications to be addressed to
_THE DIRECTOR,°
| ‘imperial Bureau of Entomology,

British Museum (Natural History),

4 Aiea aie ete London, S.W.7.

The publication and distribution of the a Bulletin of Entomological
: Research Mis has. now ieipse taken over by

THE ASSISTANT DIRECTOR, )
ago Imperial Bureau of Entomology, ‘
Bs ate 41, Queen’s Gate, London, S.W.7,

‘ ‘and Aedes and “gubscriptions should be een direot to him: or " through:
Rs tring bookseller. Dhar |

The annual ubsortion to Mol. XI | is Fifteen Panne post free.

(ee

| ‘The Editor will be pleased to receive for publication papere or
‘notes dealing with any insects that are of economic importance.
" Such communications to be addressed to

THE DIRECTOR,
| imperial Bureau of Entomology,
British Museum (Natural History),
London, S.W.7.

Orders and subscription for the ‘‘ Bulletin of Entomological Research,”
of which there are four parts in each volume, should be sent to—

THE ASSISTANT DIRECTOR,
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41, Queen’s Gate, London, S.W. 7,

or through any bookseller.

The annual subscription (in advance) to Vol. XIII is Fifteen Shillings,
- post free ; separate parts Five Shillings each, post free;

BACK VOLUMES, These may be obtained at part price, viz :—Vol. i,
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CONTENTS.

ORIGINAL ARTICLES.

Dry, " w. ‘Trypanosomiasis, in* the Absence of Tsetses,
| and a Human Disease possibly eae by ss adtonteds ds in |
Kenya Colony

EDWARDS, F. W.. A Revision of the Mosquitos of the
Palaearctic Region (illustrated).

FERRIS, G. F. Some Coccidae from hasern Asia (itustaed. )

- Lioyp, Dr. ee Notes on a Colour Tropism of Asterochiton |
(Aleurodes) vaporariorum, Westwood (illustrated).

Maccrecor, Matcorm EF, The Influence of oh ay tak alae i
Mosquito Life in Surrey:

' -MOKRZECKI, Prof, 5. A. Agrilus foveicollis, Mars., as a Cause :
>» of the Whee of the Culture of Roses in Mahe iaeis
(tllustrated), uA ae ae ahs i ea ih ae

Morris, Huspert M. The taeval and lade Stages of the a | ;
Bibionidae. Part I (illustrated). Re AAR ered 0 -

Patton, Major W. 5. Notes of the Myiass-producing ;
Diptera of Man and Animals (illustrated) . . vets

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Orders and subscription for the “ Bulletin of Entomological Research,’’

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CONTENTS.

ORIGINAL ARTICLES.

AUSTEN, Major E. E. Some Siamese Tabanidae (illustrated)

BruneEttI, E. A new Pipunculid Parasitic on Leaf-hoppers
in India (¢llustrated) ..

BRYANT, G. E. Some new injurious Phytophaga from Africa
(illustrated)

Hirt, G. F. On some Australian Termites of the Genera
Drepanotermes, Hamitermes and Leucotermes (illustrated)

James, Lt.-Col. S. P. Mosquito Life in Surrey during 1921

LAMBORN, W. A. The oo of some Ports of China
and Japan ;

Patton, Major W. S. Notes on the Species « of he nite
Musca, Linnaeus—Part I.

PAwANn, J. L. On the Eggs and HOGa nna: of Panebhors
(Janthinosoma) posticata, Wied. (Culicidae)

Pomeroy, A. W. J. New Species of African Simuliidae and
further Studies of the early Stages (zllustrated)..

RAMACHANDRA Rao, Y. Notes on the Life-histories of two
Mesopotamian Moths

SUBRAMANIAM, T. V. Some natural Enemies of Mango
Leaf-hoppers (Idiocerus spp.) in India (alustrated)

Tams, W. H. T. Description of a new ae of a
from South India: (zllustrated)

THEOBALD, F. V. A new Aphid Genus and ees found
in England (cllustrated) ay WA as

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